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Full text of "Proceedings of the United States National Museum"

SMITHSONIAN INSTITUTION 
UNITED STATES NATIONAL MUSEUM 



PROCEEDINGS 



OF THE 



UNITED STATES NATIONAL MUSEUM 



VOLUME 116 

NUMBERS 3494-3507 




UNITED STATES 

GOVERNMENT PRINTING OFFICE 

WASHINGTON : 1966 



Publications of the United States National Museum 

The scientific publications of the United States National Museum include two 
series, Proceedings of the United States National Museum and United States 
National Museum Bulletin. 

In these series are published original articles and monographs dealing with 
the collections and work of the Museum and setting forth newly acquired facts 
in the fields of anthropology, biology, geology, history, and technology. Copies 
of each pubHcation are distributed to libraries and scientific organizations and 
to specialists and others interested in the various subjects. 

The Proceedings, begun in 1878, are intended for the publication, in separate 
form, of shorter papers. These are gathered in volumes, octavo in size, with 
the publication date of each paper recorded in the table of contents of the volume. 

In the Bulletin series, the first of which was issued in 1875, appear longer, 
separate publications consisting of monographs (occasionally in several parts) 
and volumes in which are collected works on related subjects. Bulletins are 
either octavo or quarto in size, depending on the needs of the presentation. 
Since 1902 papers relating to the botanical collections of the Museum have been 
published in the Bulletin series under the heading Contributions from the United 
States National Herbarium. 

Frank A. Taylor 
Director, United States National Museum 
n 



CONTENTS 



Clarke, J. F. Gates. Neotropical Microlepidoptera, VI: 
Genera Orsotricha Meyrick and Palinorsa Meyrick (Gele- 
chiidae, Oecophoridae) . Four figures and two plates. 
No. 3502, published November 23, 1964 197-204 

New species: Palinorsa acritomorpha, P. zonaria. 

New combination: Palinorsa raptans. 

Cohen, Daniel M. A review of the Ophidioid fish genus 
Oligopus with the description of a new species from West 
Africa. Five plates. No. 3494, published October 13, 
1964 1-22 

New species: Oligopus longhursti. 
New name: Oligopiis waikiki. 

Duckworth, W. Donald. North American Stenomidae 
(Lepidoptera: Gelechioidea). Forty-five figures, four 
plates, and twelve maps. No. 3495, published October 

27, 1964 23-72 

New species: Antaeotricha fuscorectangulaia, Mothonica kimballi. 

Duckworth, W. Donald. Neotropical Microlepidoptera, 

IV: A new genus of Stenomidae with descriptions of four 

new species (Lepidoptera: Gelechioidea). Five figures. 

No. 3497, published September 11, 1964 97-114 

New genus: Lethata. 

New species: Lethata buscki, L. fusca, L. maculata, L. ruba. 
New combinations: Lethata irochalostida, L. aromatica, L. asthenopa, 
L. hoviyiella, L. glaucopa, L. invigilans, L. leucothea, L. pyrenodes, 
L. satyropa. 

Duckworth, W. Donald. North American Moths of the 
genus Swammerdamia (Lepidoptera: Yponomeutidae). 

Three figures. No. 3507, published May 25, 1965 549-556 

Gehlbach, Frederick R. Herpetology of the Zuni Moun- 
tains region, northwestern New Mexico. Ten figures and 

four plates. No. 3505, published February 26, 1965 243-332 

Nakahara, Waro. Contributions to the knowledge of the 
Hemerobiidae of western North America (Neuroptera). 
Four figures and one plate. No. 3503, published January 

26, 1965 205-222 

New species: Sympherobius brunneus, S. texanus, Kimminsia 
olympica, K. alexanderi, K. melaleuca. 

ni 



IV PROCEEDINGS OF THE NATIONAL MUSEUM 



Obraztsov, Nicholas S. Neotropical Microlepidoptera, V: 
Synopsis of the species of the genus Proeulia from Central 
Chile (Lepidoptera: Tortricidae). Nine plates. No. 

3501, published October 29, 1964 183-196 

New species: Proeulia aeihalea, P. triquetra, P. inconspicua, P. 

apospasta, P. cneca. 

New combinations: Proeulia leoninn, P. auraria, P. tenoniias, P. 

chrysopieris. 

Richards, O. W. A contribution to the study of the genus 

Sphaerocera Latreille in Central and South America 

(Diptera: Sphaeroceridae). Twenty-eight figures. No. 

3504, published April 5, 1965 223-242 

New species: Sphaerocera ecuadoria, S. trapezina, S. tertia, S. 
chimborazo, S. levicastilli, S. shannoni, S. amphora, S. 
xiphosternurn, S. musiphila, S. transversalis, B. lepida, S. trans- 
versa, S. dissecta, S. guttula. 

ScHULTz, Leonard P. Three new species of frogfishes from 
the Indian and Pacific Oceans with notes on other species 
(Family Antennariidae). Three plates. No. 3500, pub- 
lished September 1, 1964 171-182 

New species: Phrynelox lochites, Aniennarius japonicus, A. indicus. 

vScuLLEN, Herman A. Review of the genus Cerceris in 

America north of Mexico (Hymenoptera: Sphecidae). 

One hundred eighty-two figures and one plate. No. 3506, 

published May 25, 1965 333-548 

New species: Cerceris bridwelli, C. buileri, C. crandalli, C. krombeini, 
C. neahminax, C. poculum, C. cochisi, C. sandiegensis, C. rufa, 
C. banksi, C. boharti, C. macswaini, C. texana, C. wyomingensis. 
New subspecies: Cerceris echo atrata, C. finitima, citrina, C. ken- 
nicottii beali, C. vanduzeei eburnea, C. dilatata chisosensis, C. 
grandis percna, C. robertsonii bifidus, C. r. miltosus, C. insolita 
albida, C. i. atrafemori, C. aequalis bolingeri, C. a. idahoensis. 
New status: Cerceris finitima vierecki Banks, C. rufinoda crucis 
Viereck and Cockerell, C. calif ornica arno Banks, C. flavofasciata 
floridensis Banks, C. femurrubrum athene Banks, C. clypeata 
dakotensis Banks, C. c. gnarina Banks, C. c. prominens Banks, 
C. nigrescens arelate Banks. 

Springer, Victor G., and Garrick, J. A. F. A survey of 
vertebral numbers in sharks. One plate. No. 3496, 
published October 16, 1964 73-96 

Weitzman, Stanley H. One new species and two rede- 
scriptions of catfishes of the South American Callichthyid 
genus Corydoras. Six figures. No. 3498, published Oc- 

ber 13, 1964 115-126 

New species: Corydoras semiaquilus. 



CONTENTS V 

Pages 

Weitzman, Stanley H. Osteology and relationships of 
South American Characid fishes of subfamilies Lebiasininae 
and Erythrininae with special reference to subtribe Nan- 
nostomina. Ten figures. No. 3499, published October 
13, 1964 127-170 



Proceedings of 
the United States 
National Museum 




SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 



Volume 116 



1964 



Number 3494 



A REVIEW OF THE OPHIDIOID FISH GENUS OLIGOPUS 

WITH THE DESCRIPTION OF A NEW SPECIES 

FROM WEST AFRICA 



By Daniel M. Cohen ^ 



Introduction 



This paper defines the genus Oligopus, presents a discussion of 
generic synonymy, and gives diagnoses of six species, one of which is 
described herein as new to science. 

The species of Oligopus have numerous fin rays and small scales, as 
is true of many other ophidioids. Because the bases of the fin rays 
almost invariably are obscured by darkly pigmented skin, most 
counts of vertical fin elements were made from X-ray photographs. 
In addition, internal skeletal characters were observed only from 
X-ray photographs. Examination of the head pores was aided by 
the use of a compressed air jet. 

These fishes secrete a thick mucous coat that often makes it difficult 
to count scales and to observe pores. Also, the papillae that mark 



1 Director, Bureau of Commercial Fisheries Ichthyological Laboratory, Fish 
and Wildlife Service. 

1 



2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

the course of the lateral line are fragile structures which may be 
lost when the mucous coat is removed. 

Following are the definitions of some of the terms and abbreviations 
used in this paper: Predorsal: shortest distance between tip of 
snout and a vertical through origin of dorsal fin; preanal: shortest 
distance between tip of snout and a vertical through origin of anal 
fin; SL: standard length; HL: head length; upper jaw: shortest 
distance between tip of snout and posterior margin of maxillary; 
greatest depth of maxillary: depth at posterior expanded region of 
bone, but not including downward projecting maxillary spine; greatest 
head width: width of head with opercular flap compressed; D: dorsal 
fin rays ; A: anal fin rays ; caudal fin rays: only those rays articulating 
with hypural plate; vertebrae: count not including hypural; lateral 
scale rows: counted from upper angle of opercle; head canals: ter- 
minology follows Robins (1959). 

I have examined material deposited in the following institutions, 
and I am much indebted to their respective curators: Academy of 
Natural Sciences of Philadelphia (ANSP); British Museum (Natural 
History) (BMNH); Museum of Comparative Zoology at Harvard 
University (MCZ); Rhodes University, Grahamstown, South Africa 
(RU); Scripps Institution of Oceanography (SIO); Stanford Uni- 
versity, Division of Systematic Biology (SU); University of Bergen 
Zoological Museum (UBZM); University of Copenhagen Zoological 
Museum (UCZM); University of Hawaii (UH); U.S. National 
Museum (USNM); University of Puerto Rico (UP). Mr. Luis 
Rivas has furnished me with information concerning the type of 
0. claudei. Mr. and Mrs. Craig Phillips have donated a specimen 
of Stygnobrotula. Dr. Frank H. Talbot and Mr. W. I. Follett X-rayed 
the type of Eutyx diagrammus. Mr. J0rgen Nielsen X-rayed the 
type of Bythites juscus. Mr. Alwyne Wheeler X-rayed specimens 
in the British Museum. I particularly thank my colleagues Dr. 
Bruce B. Collette and Dr. Ernest A. Lachner for their advice during 
the course of this study and for their critical review of the manuscript. 

Genus Oligopus Risso 

Oligopus Risso, 1810, p. 142 (type species by monotypy Oligopus ater Risso, 1810). 
Gadopsis (not Agassiz, 1845; not Richardson, 1848) Filippi, 1856, p. 170 (type 

species by monotypy Oligopus ater Risso, 1810). 
Grammonus Gill in Goode and Bean, 1896, p. 315 (type species by monotypy 

Oligopus ater Risso, 1810). 
Verater Jordan, 1919a, p. 343 (proposed as a replacement name for Pteridium 

Filippi and Verany, 1859; however, these authors used Pteridium Scopoli, 

1777. Type species by original designation of Jordan Oligopus ater Risso). 
Eutyx Heller and Snodgrass, 1903, p. 224 (type species by monotypy Eutyx 

diagrammus Heller and Snodgrass, 1903). 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN 3 

Bathystorreus Howell Rivero, 1934, p. 69 (type species by original designation 
Benihocometes claudei Torre, 1930). 

Diagnosis. — Gill membranes separate. Chin barbel absent. 
Live-bearing. Pelvic fins each with one ray, originating close to 
level of posterior margin of preopercle and about an eye diameter 
behind symphysis of cleithra; vertical fins continuous, covered proxi- 
mally with thick, scaleless skin; pectoral fin entire, without separate, 
elongated rays. Head partly naked, bearing dermal papillae; anterior 
nostril tubular, located directly above upper lip; gill rakers 2 or 3; 
tongue a massive structure with anterior, prowlike projection fitting 
between 2 heads of geniohyoideus muscle. Branchiostegal rays 8. 
Body relatively short, stubby; head not depressed, height greater than 
^vidth. Lateral line \vith 2 or more series, dorsal and ventral some- 
times overlapping or parallel for entire lengths ; palatine lacking teeth ; 
ventrally directed projection usually present at posteroventral section 
of maxillary; posterior portion of maxillary expanded. Eyes well 
developed. Body completely covered with small cycloid scales. 
Lining of peritoneum pale in color. 

First neural spine low; neural spines 2, 3, 4 raised, followed by 
series of low neural spines with more or less truncate tops (fig. 1). 
Vertebrae 1 and 2 without ribs ; vertebra 3 with pair of antero ventrally 
directed ribs; vertebrae 4-6 with postero ventrally directed ribs 
articulating with centra; vertebra 7 with ribs attached, free or at 
end of parapophyses ; vertebrae 8-10, 11 or 12 with ribs at ends of 
parapophyses ; all other vertebrae lacking pleural ribs. 

Relationships. — Oligopus is a distinctive group of well-differentiated 
species. The genus apparently represents a rather generalized type 
of ophidioid, at least among the live-bearers, and this fact, along with 
its relatively shallow-water habitat, suggests that, together with 
Bythites, Oligopus may be close to the stem from which various 
forms descended toward an abyssal habitat while others colonized 
the coral reefs: Oligopus has affinities with fishes that dwell in both 
areas. 

Among the reef inhabitants, Microbrotula, Grammonoides, and 
Stygnobrotula are related to Oligopus; the former two even possess 
the peculiar projection on the maxillary. None of the three genera, 
however, displays the peculiar shortened and truncate neural spines 
(fig. 1) that are characteristic of Oligopus; moreover, Microbrotula 
has palatine teeth, and Bohlke (1957) has given a number of reasons 
for separating Stygnobrotula from Eutyx (here considered a junior 
synonym of Oligopus), most of which serve to separate Stygnobrotula 
from the more inclusive genus Oligopus. Particularly important are 
Bohlke's items 1, 2, 3, 4, 6, 7, 9, and 13 of his table 1, p. 3, to 



4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

which the reader is referred.^ Grammonoides is also different in 
dentition, lacking the bands of granular teeth on the premaxillary 
and the dentary. 

Among the genera confined to deeper water, the closest relatives 
of Oligojms are those forms with a tubular anterior nostril placed 
directly over the upper lip, reduced squamation on the head, and a 
single ray in each ventral fin. This group includes Diplacanthoyoma, 
Myxocephalus, the species presently assigned to Cataetyx (at least 
two genera are included in this group), and Bythites. All of these 
differ from Oligopus in possessing palatine teeth, and in lacking 
shortened, truncate neural spines. In addition, Diplacanthopoma 
and Myxocephalus lack scales on the head. Most of the species of 
Cataetyx (though not the type, C. messieri) have strongly depressed 
heads. 

Oligopus ater, the type species of Oligopus, long has been considered 
congeneric with a superficially similar species that was described 
originally by Doderlein (1886) as Pteridium armatum. Bougis and 
Ruivo (1954) reported on specimens that they identified as Ben- 
thocometes robustus, a name that they treated as a senior synonym of 
P. armatum. They illustrated (their fig. 19) specializations of the 
anterior vertebrae that bear some resemblance to those long known 
to exist in the Ophidiidae (in the restricted sense), but there is some 
conflictin this identification, for Doderlein (1886) did not show modified 
anterior vertebrae in his figure of a dissection of Pteridium armatum. 
These structures, however, have been shown to be sexually dimorphic 
in the ophidiid genus Ophidion (Rose, 1961). Whatever the identity 
of P. armatum, Oligopus differs widely both from Doderlein's descrip- 
tion and from Benthoco metes. It is of interest to note that another 
species of Oligopus (0. claudei) was also confused with Benthocometes. 

The foregoing discussion is based on the material listed below. 

Material examined. — Benthocometes robustus (USNM 29057, 
paratype) ; Bythites juscus (UCZM, holotype) ; Bythites lepidogenys 
(USNM 74152, holotype); Cataetyx hawaiiensis (USNM 162715, 
holotype) ; Cataetyx laticeps (UBZM, holotype) ; Cataetyx messieri 
(BMNH, holotype) ; Diplacanthopoma brachysoma (BMNH, holotype) ; 
Diplacanthopoma brunnea (USNM 74148, holotype); Grammonoides 
opisthodon (RU, holotype); Microbrofula rubra (USNM 162710, 
holotype); Myxocephalus japonicu^ (USNM 160604); Stygnobrotula 
latebricola (USNM 187777); Xenobythites armiger (USNM 74153, 
holotype). 

Synonymy. — In his original description of Oligopus ater, Risso 
(1810) credited the genus Oligopus to Lacepede; however, Risso's 

2 Boeseman (1960) overlooked the description of Stygnobrotula latebricola and 
described the same species as Eutyx tumidirostris. 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS COHEN 5 

spelling is an unjustified emendation of Oligopodus Lacepede, 1800, 
proposed for Conjpliaena velifera Pallas, and therefore, Oligojpus is 
available from 1810 with Risso as the author. 

Swainson (1839) placed 0. ater in Pteridium Scopoli, and Filippi (1856) 
proposed the generic name Gadopsis (already twice preoccupied) for 
the species. Filippi and Verany (1859) then accepted Pteridium 
ScopoU (proposed for Coryphaena velifera Pallas) for 0. ater and have 
been followed by numerous authors. Gill {in Goode and Bean, 1896) 
proposed Grammonus for Oligopus ater Risso, with no comment. 

Verater was first proposed by Jordan (1919a) as a substitute for 
Pteridium Filippi and Verany although these authors plainly indicated 
they were following Swainson, who used Pteridium Scopoli; however, in 
a later publication during the same year, Jordan (1919b) presented 
Verater as a replacement name for Gadopsis Filippi, with 0. ater 
Risso as the type species. 

Heller and Snodgrass compared Eutyx with Grammonus Gill and 
separated the two on the grounds that the former genus lacked an 
opercular spine, had a double lateral line, and had large muciferous 
canal openings on the head. Actually, Eutyx may have its small 
opercular spine hidden or exposed (Bohlke, 1957, and the present 
author) ; its lateral line (described below under Oligopus diagrammus) 
is similar to that in other species of the genus, and all of the species 
here referred to Oligopus have canal openings on the head. 

Bathystorreus was proposed for a species originally described in 
Benthocometes , where it obviously did not belong. The single known 
specimen is in bad condition ; however, an X-ray photograph showing 
the abdominal vertebrae gives reason enough to place this species 
in Oligopus. 

Species. — 'Six species are referred herein to Oligoyus. Although 
the available material of most is limited, a few suggestions concerning 
relationships and distributions can be presented. 0. claudei from 
the tropical western Atlantic is distinct from all other Oligopus. 
Distinct preopercular spines and numerous head pores are its distin- 
guishing characters. 0. diagrammus is known from Galapagos, 
Guadalupe, and lower Baja California. The material suggests the 
presence of a species complex or a group of subspecies. 0. diagrammus 
has a reduced number of head pores but high fin ray and vertebral 
counts. Apparently it is most distantly related phylogenetically 
to the western Atlantic species. The other species are 0. longhursti 
(herein described as new), known from tropical West Africa; 0. ater 
from the Mediterranean; 0. robustus from Japan, the Philippines, 
and the Indian Ocean; and 0. waikiki from Hawaii. The latter, wide- 
ranging group of four species is intermediate phylogenetically as 



6 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

well as geographically between the western Atlantic and the eastern 
Pacific species. 

I do not consider Grammonus leucos Osorio, 1917, to be referable 
to Oligopus, as the original description mentions the presence of teeth 
on the palatines. Professor Fernando Frade has informed me that 
the type cannot be found in the Museu Bocage. 

Key to Species of Oligopus 

la. One or no pores in lateral head canal system; 1 or 2 pores in supraorbital 
canal. Spines along preopercle margin absent or small and weak. 
2a. Dorsal fin rays 68-87; anal fin rays 51-62; vertebrae 40-46. 

3a. Dorsal fin rays 68-74; anal fin rays 51-52; vertebrae 40-42 . . . O. ater 
3b. Dorsal fin rays 83-87; anal fin rays 57-62; vertebrae 44-46. 

4a. Lateral scale rows about 120. Upper row of lateral line organs 
about 35-45; lower about 35-40; dip in ventral lateral line in vent 

area O. longhursti 

4b. Lateral scale rows about 75-85. Upper row of lateral line organs 
about 21; lower row about 25-30; no dip in ventral lateral line. 

O. robustus 
2b. Dorsal fin rays 93-115; anal fin rays 71-91; vertebrae 48-53. 

5a. Lateral scale rows about 80; anal fin rays 71; 6 pores in infraorbital 

canal system O. waikiki 

5b. Lateral scale rows 97-115; anal fin rays 76-91; 5 or fewer pores in infra- 
orbital canal system O. diagrammus 

lb. Three or 4 pores in lateral head canal system; 2, 3, or 4 pores in supraorbital 
canal. Several small but distinct spines along preopercle margin. 

O. claudei 

Oligopus ater Risso 

Plate 2 

Oligopus ater Risso, 1810, p. 142, pi, 11, fig. 41 (original description. Gulf of Saint 
Hospice, France). — Fowler, 1936, p. 1329 (description, synonymy). 

Oligopiis niger Risso, 1826, p. 338 (new name for 0. ater Risso, 1810; description). 

Pteridium ater Swainson, 1839, p. 302 (0. ater Risso listed under Pteridium Scopoli). 

Gadopsis ater Filippi, 1856, p. 170 (description of swim bladder, new genus pro- 
posed) . 

Pteridium alrimi Filippi and Verany, 1859, p. 195, fig. 6 (synonymy, description, 
swim bladder, relationships). — Doderlein, 1886, p. 73 (comparison with 
Pteridium armatxim). — Giinther, 1887, p. 105 (description, 1 specimen from 
Mediterranean). — Bellotti, 1888, p. 222 (sexual dimorphism in dentition of 
specimens from Nice and nearby, comparison with Pteridium armatum.) — 
Lo Bianco, 1909, p. 741 (pelagic postlarvae circa 30 mm., January, February, 
March, in Gulf of Naples). — D'Ancona, 1938, p. 159, figs. 2, 4, 8 (comparison 
with P. armatum, description, based on material from Nice, Naples, and 
Zirona, nomenclature, relationships, distribution, additional references not 
here cited). 

Grammonus ater Goode and Bean, 1896, p. 317 (new genus proposed, synonymy, 
compiled description). — Tortonese, 1958, p. 333 (listed, doubts that G. ater 
(Risso) and G. armatus (Doderlein) belong in different genera). 

Verater ater Jordan, 1919a, p. 343 (new genus proposed). 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN 7 

Misidentifications: Pteridium atrum Emery, 1885, p. 158, fig. 21 (30 mm. specimen, 
description, probably Benthocometes robustus). — Roule and Angel, 1930, 
p. 110, pi. 6, fig. 146 (Azores, description of postlarvae). 

Study materials. — 1 specimen, Nice, MCZ 26457. 

Counts and measurements.— (Measurements given in mm., 
followed by percent of standard length in parentheses) D 74; A 52; 
pectoral 19; vertebrae 40; caudal 10; vertical scale rows about 83; 
SL 89; body depth at dorsal origin 17.5 (19.7); predorsal 31.2 (35.0); 
preanal 50.0 (56.1) ;HL 25.0 (28.0); snout 5.8 (6.5); orbit 3.0 (3.4); 
upper jaw 14.2 (15.9); greatest maxillary width 4.5 (5.1); greatest 
head width 11.0 (12.3). (See D'Ancona, 1938, p. 162, for additional 
data.) 

Description. — Squamation: Scales present on large area on top of 
head behind eye level; present on side of head on opercle and in area 
ahead of opercle extending forward nearly to eye and ventrally to 
region behind maxillary. Dorsal and lateral scale patches separated 
by scaleless lateral canal. 

Lateralis system: Lateral canal with single pore near upper angle of 
opercle. Supraorbital canal with 2 pores: 1 in front of, and below, 
anterior nostril; the other, difficult to find, above anterior nostril. 
Infraorbital canal with 6 pores: 3 beneath nostrils in skinfold over 
upper jaw; 1 very small pore close to level of posterior part of eye; 2 
larger pores above posterior expanded part of maxillary. Preoper- 
culomandibular canal with 8 pores: 2 at tip of dentary, 1 opening 
anteriorly, the other posteriorly; 1 beneath lower lip at about midlength 
of snout; 1 at about midlength of jaw; 1 shghtly ahead of posterior 
margin of maxiUary; 1 slightly behind same; 2 on posterior margin of 
preopercle. 

Circumorbital and interorbital areas and side of snout cavernous, 
covered with thick skin bearing scattered papillae. Interorbital area 
particularly rugose. 

Lateral line marked by small, dark, dermal filaments. Series of 23 
originates above opercle. Between opercle and level of dorsal fin 
origin filaments in elongate, irregular cluster from which lateral line 
extending posteriorly in straight line, between midline and dorsal 
profile. Dorsal line terminating 63 mm. from tip of snout. Second 
line of about 40 filaments originating in midline close to level of pos- 
terior tip of pectoral fin, descending, then rising to midline of body to 
form shallow irregular semicircle with lowest point opposite vent, 
then extending straight back in midline of body to tail. Filaments 
similar to those along lateral lines scattered about on head. 

Dentition: Premaxillary with band of uniform granular teeth. 
Dentaries with similar teeth and irregular row of larger, conical 
teeth along inner edge of granular band. Head of vomer a broad 



8 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

V with arms expanded. Both short granular and larger conical teeth 
present on vomer. According to Bellotti (1888), females of this 
species have the type of dentition noted above, although without 
larger teeth on the vomer, while males have both larger teeth and 
granular teeth on the premaxillary, vomer, and dentary. 

Head spines: Short, sharp-pointed spine piercing skin at upper 
corner of opercle. Posteroventral margin of preopercle with 2 
blunt projections at angles. Ventrally directed spine at end of 
maxUlary prominent, piercing skin. 

Vertebrae and ribs: Neural spines 5-13 short and broad but be- 
coming progressively more elongate and narrower, their tips truncate; 
neural spines on subsequent centra needle-like. Centra 7-11 with 
ribs at ends of parapophyses, subsequent centra lacking pleural ribs. 
Centra 4-8 with epipleurals attached to pleural ribs; on centra 9-12 
epipleurals appear associated with parapophyses. 

Color: Body light brown, vertical fins and head darker. Long 
in preservative, this specimen evidently was much darker in life. 
Risso used the conmion name "fanfre negre" and stated that the 
species was black. 

Distribution. — KJiown from the area around Nice, from Naples, 
and from several locaUties in the Adriatic. Little is known of its 
depth distribution. 

Habitat. — Dwells in rocky areas in caves, according to Risso 
(1810), and apparently is secretive. 

Comments. — Professor D'Ancona (1938) has presented an excellent 
review of this species, and his paper should be consulted for informa- 
tion and references not given here. 

Oligopiis longhursti, new species 

Plate 3 

Study material. — All females. Holotype: USNM 187778, off 
Lagos, Nigeria in 5-10 fathoms, otter trawl, collected by A.R. Long- 
hurst, August 1961. Paratypes: USNM 187779, 1 specimen, data 
as for holotype; USNM 191732, 2 specimens, Monrovia, Liberia, 
Bush Rod Island, beach seine, collected by George C. Miller, October 
7, 1952; USNM 193678, 1 specimen, Liberia, 3-7 fathoms off mouth 
of St. Paul River, trawl, collected by George C. Miller, October 14, 
1953. 

Counts and measurements. — See table 1. 

Description. — Squamation: Scales present on large area on top 
of head behind eye level; present on side of head over opercle and 
area in front of dorsal half of opercle, which extends to within eye 
diameter of eye. Lateral scale patch bounded dorsally by scaleless 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN 9 

lateral canal and ventrally by scaleless area that demarcates separate 
patch of scales posterior to rear margin of maxillary. 

Lateralis system: Lateral canal with 1 pore near upper angle of 
opercle. Supraorbital canal with 2 pores: 1 in front of anterior 
nostril; the other, very small, above anterior nostril. Infraorbital 
canal with 5 or 6 pores: 3 in row behind anterior nostril; 1, very 
small, near level of posterior margin of eye (absent in USNM 193678) ; 
and 2 above posterior expanded portion of maxillary. Preoperculo- 
mandibular canal with 8 pores: 2 at tip of dentary, 1 opening 
anteriorly, the other posteriorly; 1 beneath lower lip close to level 
of posterior nostril; 1 slightly ahead of level of posterior margin of 
maxillary; 1 near lower angle of same; 2 shaped like elongate slits, on 
posterior margin of preopercle. 

Cu'cumorbital and interorbital areas and side of snout cavernous 
and covered with thick skin bearing many small, dark, dermal papillae 
similar to those marking course of lateral lines along side of body. 
These filaments particularly abundant in interorbital region. 

One series of lateral line filaments numbering about 35-45 and 
originating on head above opercle and, after slight irregular dip 
near origin, extending posteriorly in straight line between midline of 
dorsal profile, this dorsal line terminating more than half way back 
along body. Second line of about 35-40 filaments originating at 
about midhne close to level of posterior tip of pectoral fin, descending, 
then rising to midline of body to form irregular semicircle having 
lowest point opposite origin of anal fin, then extending in straight 
line in midline of body to tail. 

Dentition: Premaxillary bearing uniform band of granular teeth. 
Dentaries bearing similar teeth plus irregular row of larger, conical 
teeth along inner edge of granular band. Head of vomer with widely 
spread wings bearing short granular teeth. All four specimens 
females. 

Head spines: Short, weak spine present at upper corner of opercle. 
May be buried beneath skin and invisible without dissection, or barely 
visible at its tip. Posteroventral margin of preopercle may be com- 
pletely rounded or show 2 slight angles. Maxillary spine varying from 
prominent projection pictured in holotype to virtually absent condi- 
tion in 1 Liberian paratype. 

Vertebrae and ribs: Neural spines 5-13 short and broad but becoming 
progressively more elongate and narrower, their tips more or less (5 
and 13) truncate; neural spines on subsequent centra needle-like. 
Centra 7-12 with ribs at ends of parapophyses, subsequent centra 
lacking pleural ribs. Centra 4-9 with epipleurals attached to pleural 
ribs; on centra 10-13 epipleurals appearing to be associated with 
parapophyses. 

724-329—64 2 



10 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Color: Head and body brown, vertical fins black. Beneath super- 
ficial brown pigmentation of scales and scale pockets large, dark 
chromatophores distributed regularly on body. 

Distribution. — Known only from the type localities. Taken in a 
beach seine and at 3-10 fathoms. 

Comments. — The holotype extruded fertilized eggs that must have 
been close to hatching and that are about 0.9 mm. in diameter. The 
Nigerian paratype has paired ovaries about 25 mm. long. Each 
contains an estimated 3,000 unfertilized eggs (based on counts and 
volumetric displacement on }^ of 1 ovary), which are about 0.5 mm. 
in diameter. Eggs are of about the same size in all parts of the ovary; 
however, the eggs closest to the posterior end are more densely granu- 
lar. One Liberian paratype (USNM 191732) extruded granular 
(unfertilized) eggs of about 0.5 mm. in diameter. The other Liberian 
specimen has its ovaries packed with an estimated several thousand 
young fish. The embryos are between 4 and 5 mm. long and have 
small yolk sacs and open choroid fissures. Elongate processes are 
attached to the embryo a short distance behind the yolk sac. 

Oligopus robiistus (Smith and Radcliffe) 

Grammomts robustus Smith and Radcliffe in Radcliffe, 1913, p. 168, pi. 13, fig. 4 
(original description, "Albatross" St. 5409, 10°38' N., 124°13' E.— Norman, 
1939, p. 79 (Gulf of Aden, 457-549 m.) .— Kamohara, 1954, p. 5, fig. 3 (5 
specimens from Kochi market, description). 

Bythites lepidogenys (not Smith and Radcliffe, 1913) Kamohara, 1952, p. 93 (3 
specimens from Mimase, description). 

Study material. — Holotype, USNM 74149, male. 

Counts amd measurements. — (Measurements given in mm., 
followed by percent of standard length in parentheses) D 87; A 62 
pectoral 25; vertebrae 46; caudal 8; vertical scale rows about 75 
SL 129; body depth at D origin 32.5 (25.2); predorsal 42.2 (32.7) 
preanal 71.1 (55.1); HL 40.4 (31.3); snout 6.5 (5.0); orbit 6.5 (5.0) 
upper jaw 22.0 (17.1); greatest maxillary width 8.2 (6.4); greatest 
head width 21.2 (16.4). Additional meristic and morphometric data 
are presented by Kamohara (1954). 

Description. — Squamation: Patch of scales present on nape. 
On sides of head, scales present on opercle and on area in front of 
opercle extending forward nearly to eye, dorsally to level of top of 
opercle, where scaleless lateral canal separates dorsal and lateral scale 
patches, and ventrally to area behind maxillary. 

Lateralis system: Lateral canal with 1 pore near upper angle of 
opercle. Supraorbital canal with 2 pores: 1 below, and in front of, 
anterior nostril tube; the other immediately above tube. Infra- 
orbital canal with 6 pores: 3 beneath nostrils over upper lip; 1 at 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN H 

level of posterior part of eye; 2 above posterior expanded portion 
of maxillary. Preoperculomandibular canal with 8 pores: 2 at tip 
of dentary, 1 opening anteriorly, the other posteriorly; 1 beneath 
lower lip near level of posterior nostril; 1 close to midlength of jaw; 
1 ahead of posterior margin of maxillary; 1 behind same; 2 at pos- 
teroventral angle of preopercle. 

Small, dark papillae scattered on head, concentrated in inter- 
orbital area. Lateral line marked by papillae similar to those on 
head. One series of 21 originating above opercle and, after slight 
irregular dip near origin, extending posteriorly in straight line be- 
tween midline and dorsal profile, terminating at level of vent. Second 
line of about 25 filaments originating in midline at level of origin of 
anal fin and extending posteriorly in midline of body to point about 
10 mm. from tail (the tail has become somewhat abraded and in all 
likelihood the lateral line actually continues to the base of the tail 
as shown by Kadcliffe, 1913, and Kamohara, 1954.) 

Dentition: Premaxillaries, dentaries, and broadly V-shaped vomer 
bearing bands of uniformly smaU, granular teeth. 

Sex: Specimen male with prominent, paired testes about 20 mm. 
long. 

Head spines: Small, sharp-pointed spine piercing skin at upper 
angle of opercle. Post ero ventral angles of preopercle with 2 very 
blunt points. Maxillary spine prominent on right side of specimen, 
weakly developed on left side. 

Vertebrae and ribs: Neural spines 5-15 short and broad, but be- 
coming progressively more elongate and narrower; their tips truncate; 
neural spines on subsequent centra more needle-like. Centra 7-11 
with ribs at ends of parapophyses, subsequent centra lacking pleural 
ribs. Centra 4-7 with epipleurals attached to pleural ribs; on centra 
8-15 epipleurals appearing to be associated with parapophyses. 

Color (in alcohol, quoted from original description): "Wood-brown; 
dorsal and anal dusky anteriorly, becoming dark clove brown pos- 
teriorly; the basal portion of caudal clove brown, distal portion some- 
what lighter; pectorals dusky; peritoneum silvery gray." 

Distribution. — Known from Mimase, Japan; the Philippines 
between Cebu and Leyte, at a depth of 189 fathoms on a green mud 
bottom; and the Gulf of Aden between 25 and 30 fathoms. 

Oligopus waikiki, new name 

Plate 4 

Microbrotula nigra Gosline, 1953, p. 220, fig. Id (original description, off Waikiki 
Reef, Oahu, Hawaii).^ 

3 If Microbrotula nigra Gosline is referred to Oligopus, the specific name must 
be rejected as a junior secondary homonym of Oligopus niger Risso, which is a 
junior synonym of Oligopus ater Risso. 



12 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii8 

Study material. — Holotype of Microbrotula nigra, USNM 179898, 
formerly UH 1684, female. 

Counts and measurements. — (Measurements given in mm., followed 
by percent of standard length in parentheses) D 98 ; A 71 ; pectoral 26 ; 
vertebrae 48; caudal 8; vertical scale rows about 72; SL 61.4; body 
depth at D origin 17 (27.7) ; predorsal 16.8 (27.4) ; preanal 30.0 (48.9) ; 
HL 15.9 (25.9); snout 3.2 (5.2); orbit 3.0 (4.8); upper jaw 9.0 (14.7); 
greatest maxillary width 3.6 (5.9); greatest head width 9.5 (15.5). 

Description. — Squamation: Patch of scales present on nape. 
Second patch covers opercle and dorsal half of area in front of it, 
extending forward to within eye diameter of eye. Small, separate 
patch consisting of few scales present behind rear margin of maxillary. 

Lateralis system: Lateral canal lacking pores. Supraorbital canal 
with 2 pores: 1 below, and in front of, anterior nostril tube; the other, 
very small, above tube. Infraorbital canal with 6 pores: 3 beneath 
nostrils over upper lip; 1 at level of posterior part of eye; 2 above 
posterior expanded portion of maxillary. Preoperculomandibular 
canal with 8 pores: 2 at tip of dentary, 1 opening anteriorly, the other 
posteriorly; 1 beneath lower lip near level of posterior nostril; 1 close 
to midlength of jaw; 1 slightly ahead of level of posterior margin of 
maxillary ; and 3 shaped like elongate slits along posteroventral and 
posterior margin of preopercle. 

Small, dark papillae scattered on muzzle and interorbital region. 
Lateral line marked by papillae. One series of about 20 originating 
above opercle and, after dip near origin, extending posteriorly to level 
of vent. Second line of about 35 papillae originating in midline at 
level of origin of anal fin and extending posteriorly in midline of body 
to tail, although posterior papillae very small. 

Dentition: Premaxillaries and dentaries bearing bands of uniformly 
small, granular teeth. Vomer with 2 patches of similar teeth. Al- 
though Gosline (1953) noted the presence of palatine teeth, I find no 
trace of them. 

Sex: Specimen female with small embryos in ovaries. 

Head spines: Small, sharp-pointed spine piercing skin at upper 
angle of opercle. Posteroventral angles of preopercle bearing several 
weakly spinous points. Maxillary spine prominently developed. 

Vertebrae and ribs: Neural spines 5-14 short and broad, but be- 
coming progressively more elongate and narrower, their tips truncate; 
neural spines on subsequent centra more needle-like. Centra 7-11 
with ribs at ends of parapophyses, subsequent centra lacking pleural 
ribs. Centra 4-8 with epipleurals attached to pleural ribs ; on centra 
9-12 epipleurals appearing to be associated with parapophyses. 

Color: Body brown, fins darker. 

Distribution. — Known only from the type locality at about 30 feet. 



PROC U.S. NAT. MUS. VOL 116 



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REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN 13 

Oligopus diagrammus (Heller and Snodgrass) 

PLATES 1, 5 

Eutyx diagrammus Heller and Snodgrass, 1903, p. 224, pi. 19 (original description, 
Galapagos, Tagus Cove, Albemarle Island, and Seymour Islands). — Bohlke, 
1957, p. 3 (paratype of E. diagrammus compared with Stygnohrotula late- 
hricola) . 

Study material. — Holotype, SU 6346 and paratype SU 7059 from 
the Galapagos. The following from Guadalupe Island, Baja Cali- 
fornia, Mexico: SIC 58-497 (1); SIC 58-493 (1); SIO 54-213A (1); 
SIO 54-219 (1); SIO H53-169 (1); SIO 54-219A (6); SIO 60-14-61E 
(2). Following from peninsular Baja California or closely adjacent 
islands: SIO 61-272-61B, Espiritu Santo Island (3) ; SIO 61-227-61A, 
Cape San Lucas (2); SIO 61-242-61 A (4); and SIO 61-249 (1), 
Los Frailes. 

Counts and measurements. — Caudal 10; vertical scale rows be- 
tween 100 and 115, very difficult to count. Vertebrae 49-53. (See 
table 2 for a summary of measurements, and table 3 for counts.) 

Description. — Squamation: Patch of scales on top of head pos- 
terior to interorbital. Patch of scales present on opercle and in front 
of opercle extending anteriorly to a point less than eye diameter 
behind eye. Dorsal and lateral scale patches appearing continuous 
in some specimens and separated by scaleless area over lateral canal 
in others (small size of scales and adherent mucous coating on head 
make this a difficult character of which to be certain). Scale patch 
posterior to hind margin of maxillary appearing to be continuously 
connected with main opercular scale patch. 

Lateralis system: Lateral canal with 1 pore near upper angle of 
opercle. Supraorbital canal with 1 pore in front of, and below, 
anterior nostril. Infraorbital canal with 5 pores: 1 behind anterior 
nostril; 2 along lip below posterior nostril; and 2 above posterior 
expanded portion of maxillary. Preoperculomandibular canal with 
8 pores: 2 at tip of dentary, 1 opening anteriorly, the other posteriorly ; 
1 beneath lower lip close to level of posterior nostril (not shown in 
fig. 4) ; 1 at about midlength of jaw (could not be found in holotype) ; 
1 slightly ahead of posterior margin of maxillary; 1 slightly behind 
same. Two elongate slits along posterior margin of preopercle. 

Circumorbital, interorbital, and opercular areas and snout bearing 
scattered papiUae. Small dark papillae distributed on head but 
thickly dispersed in particular on top of head and over adjoining 
predorsal area of top of body. 

Lateral line marked by small papillae. One series of about 30 
originating on head above opercle and, after slight irregular dip 
behind opercle, extending posteriorly in straight line between midline 



14 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

and dorsal profile. Dorsal line terminating slightly more than half 
way back along body. Second line of about 37-45 small papillae 
originating in midline close to level of posterior tip of pectoral fin, 
not making pronounced dip at level of vent but extending straight 
back to tail with, at most, a slight irregularity near its origin. 

Dentition: Dentaries with irregular row of sharply pointed teeth 
along inner margin of bone. Tooth-bearing area of bone broadens 
on anterior K to % of lower jaw. In specimens of less than about 
100 mm., area exterior to large teeth occupied by granular teeth. In 
larger specimens these show gradual transition to second row of 
larger conical teeth that parallels inner row. Also 2 types of dentition 
on preraaxillary. Specimens from 34 mm. to 81 mm. with uniform 
band of granular teeth, while those measuring 81 mm. and larger 
bearing row of large conical teeth along outer edge of band of granular 
teeth. Smaller specimens with only small granular teeth in broadly 
V-shaped patch on head of vomer. Larger specimens with larger 
conical teeth as well. Dentition in this species does not appear to 
be influenced by sex. 

Sex: Two types of external sex organs, one consisting of stiff hood 
projecting posteriorly from vent and folding over genital area, the other 
consisting of only a low, fleshy ridge in place of prominent hood. In 
either type a delicate white papilla of varying length may be evident. 
Neither the two types of external genitalia nor the occurrence of the 
papilla appears to be correlated with sex as determined by gross 
observation of the gonads. In addition, at least one specimen (SIO 
61-272-61B, Espiritu Santo) and possibly others appear hermaphro- 
ditic. What I have interpreted as testicular tissue produces sper- 
matophore-like objects rather than free-running sperm suspensions. 
If my observations are correct, then sex in this species is a complex 
problem that requires detailed study. 

Head spines: Blunt, fiat spine deeply buried in flesh of opercle in 
most specimens, but exposed in paratype. Ventrally directed maxil- 
lary spine prominent in most specimens. 

Vertebrae and ribs: Neural spines 5-14 short and broad but be- 
coming progressively more elongate and narrower, their tips truncate; 
neural spines on subsequent centra needle-like. Centra 7-11 with 
ribs at ends of parapophyses (although in some, ribs appear to ar- 
ticulate with centrum no. 7) ; subsequent centra lacking pleural ribs. 
Centra 4-9 with epipleurals attached to pleural ribs; 10, 13, 14, or 15 
with epipleurals associated with parapophyses or haemal spines. 

Color: Brown in preservative. In life: "Dark brown, head pur- 
plish-brown; fins blackish" (Heller and Snodgrass,1903). 

Distribution, — Previously known only from the Galapagos at 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN 15 

about 3 fathoms. Here recorded from Guadalupe Island and Baja 
California. 

Comments. — The data on dorsal and anal fin ray counts presented 
in table 3 indicate that the Galapagos-Guadalupe populations and 
the Baja California populations of 0. diagmmmus have differentiated. 
Under ordinary circumstances I would recognize them as named taxa ; 
however, it seems likely that additional populations will be discovered 
in the tropical eastern Pacific, and this, combined with the puzzling 
nature of sexuality in this species, makes me hesitant to do more than 
point out the differences and the problems involved in interpreting 
them. 

Oligopus claudei (Torre) 

Benthocometes claudei Torre y Huerta, 1930, opposite p. 171, unnumbered plate 
(original description, Cuba, Matanzas Bay) ; 1931?, p. 231, fig. (brief dis- 
cussion) . 

Bathyslorreus claudei Howell Rivero, 1934, pp. 69-72, pi. 7 (new genus proposed, 
description). 

Grammomis mowbrayi Grey, 1951, p. 154, fig. 1 (original description, type locality, 
Bermuda). — CoUette, 1962, p. 443 (Bahamas). 

Study material. — The holotype of B. claudei, MCZ 33943 (in very 
poor condition) . Three specimens ANSP Chaplin Bahama collections, 
St. 513; about Yi mile N. of the center of Green Key, Bahamas; taken 
from inside a small, isolated coral head at a depth of 50 feet. One 
specimen, UP 1150; Puerto Rico, La Parquera, 6 miles offshore at 
vertical ledge, 65 feet. One specimen, UP 1151, Curasao, cave at 
shore, about 100 meters SE. of entrance to bay. 

Counts and measurements. — See table 4. 

Description. — Squamation: Scales present in patch over nape, on 
side of head over opercle and cheek, extending forward to a point less 
than eye diameter behind eye and ventrally to patch behind posterior 
margin of maxillary. 

Lateralis system: Lateral canal with 4 pores in row: the most posterior 
above upper angle of opercle ; the most anterior at junction of lateral canal 
and infraorbital canal. Supraorbital canal in Bahaman specimens 
with 4 pores extending in straight line from most anterior in front of 
anterior nostril to most posterior over rear half of eye; Puerto Rican 
specimen with only the 2 most anterior pores; and Curagao specimen 
with the 3 most anterior pores on one side and the 2 most anterior pores on 
other side. Infraorbital canal with 8, 9, or 10 pores: 3 beneath nos- 
trils; 1 smaller pore beneath eye about midway between orbit and lip; 
2 or 3 pores in skin along maxillary sheath; and 2 or 3 pores in row 
extending up to lateral canal. Preoperculomandibular canal with 
9-11 pores: 2 at tip of dentary, 1 opening anteriorly, the other posteri- 
orly; 1 beneath lower lip at level of anterior margin of eye; row of 3 



16 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

pores along medial edge of dentaiy, anteriormost about K of distance 
along pore, posterior beneath end of maxillary: 1 small pore sometimes 
present lateral to middle pore of this series; 1 pore above angular; 2 
on posterior margin of preopercle, and sometimes another above, and 
in front of, dorsalmost preopercular pore. The pore system is well 
illustrated in the figure given with the original description of this 
species (Torre y Huerta, 1930). One difference is that the Bahaman 
material lacks the pore on the posterior surface of the angular, as 
shown on the holotype. 

Lateral line system along body variable and complex, consisting 
basically of 2 rows of papillae: one with about 35 papillae originating 
over opercle and extending posteriorly almost to tail in line between 
midline of fish and dorsal profile; the other, with about 25 papillae, 
beginning slightly ahead of level of vent and, after an irregular be- 
ginning, proceeding posteriorly almost to tail in straight line between 
midline of fish and ventral profile of fish. In addition to these 2 
basic lines (illustrated on the figure of the holotype), there are a 
profusion of other papillae whose distribution will not be described 
in detail. Suffice it to say that some are distributed virtually at 
random, some are in short series above or below the main lines, some 
are in short rows in the midline of the fish. 

Papillae also scattered about on head, as are very small dermal 
filaments; however, no rugose areas on head. 

Dentition and sex: In two specimens a band of granular teeth on 
premaxillary, another on dentary. Head of vomer carrying 2 large 
bony knobs, each with patch of granular teeth, each patch connected 
by narrow line of short teeth. Three other specimens with bands 
of granular teeth noted above and, in addition, with irregular 
single row of larger conical teeth located lateral to granular row on 
premaxillaries and medial to granular row on dentaries. A few of 
vomerine teeth also slightly enlarged. Specimens with granular teeth 
with only a small, fleshy flap over genital area. Other 3 with promi- 
nent, stiffened hoods; at least 1, UP 1151, with well-developed testes. 

Head spines: Short, sharp spine piercing skin at upper angle of 
opercle. One specimen with several short spines at lower angle of 
opercle. Two short, sharp spines at lower angle of preopercle. 
Ventrally projecting premaxillary spine prominent, angular project- 
ing downward as prominent blunt spine. 

Vertebrae and ribs: Neural spines 5-13 or 14 short and broad but 
becoming progressively more elongate and narrower; their tips trun- 
cate. Neural spines on subsequent centra needle-like. Centrum 
7 with free ribs, although rudiments of parapophysis visible on X-ray 
photograph of 1 specimen. Centra 8, 9, and 10-12 with ribs at ends 
of parapophyses, subsequent centra lacking pleural ribs. Centra 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN 17 

4-8 with epipleurals attached to pleural ribs; on centra 9-13 or 14, 
epipleurals associated with parapophyses or haemal spines. 

Color: Body light brown; vertical fins dark, particularly on their 
vertical margins. 

Distribution. — Bermuda, the Bahamas, Puerto Rico, and 
Curagao. 

Comments. — The holotype of 0. claudei is in such poor condition 
that accurate measurements are nnpossible, and observations on the 
lateralis system are subject to question. It does seem certain, how- 
ever, that, with the material at hand, there is no way of maintaining 
Grammonus mowhrayi Grey as a valid species. 

Literature Cited 

Bellotti, Cristoforo 

1888. Notte ittiologiche, X: Pteridium atrurn Risso. Atti. Soc. Italiana 
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BOESEMAN, M. 

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BoHLKE, James E. 

1957. A new shallow-water brotulid fish from the Great Bahama Bank. 
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44, pp. 135-176, pis. 7-117. 
Risso, A. 

1810. Ichthyologie de Nice, etc. Paris, xxii-|-338 pp., 11 pis. 
1826. Histoire des principales productions de I'Europe M6ridionale et 
particuh^rement de ceUes des environs de Nice et des Alpes Mari- 
times, vol. 3, xvi-|-480 pp., 16 pis. 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS COHEN 19 

Robins, C. Richard 

1959. Studies on fishes of the family Ophidiidae, III: A new species of 
Lepophidium from Barbadoes. Breviora, Mus. Comp. Zool., no. 
104, 7 pp. 
RouLE, Louis, and Angel, Fernand 

1930. Larves et alevins de poissons provenant des croisieres du Prince 
Albert I" de Monaco. R6s. Camp. Sci. Albert I", fasc. 79, 139 
pp., 6 pis. 
Rose, Jo Ann 

1961. Anatomy and sexual dimorphism of the swim bladder and vertebral 
column in Ophidion holbrooki (Pisces: Ophidiidae). Bull. Mar. 
Sci. Gulf and Caribbean, vol. 2, no. 2, pp. 280-308. 
SwAiNsoN, William 

1839. The natural history of fishes, amphibians and reptiles or monocar- 
dian animals, vol. 2, vi + 452 pp. 
Torre y Huerta, Carlos de la 

1930. Un pez de nueva especie, el "Benthocometes claudei Torre" cogido 
en el tubo de la planta de M. Claude. Rev. Soc. Geogrdf. Cuba, 
aiio 3, num. 4 [unnumbered page inserted between pp. 170-171]. 
1931? Un pez de nueva especie, el "Benthocometes Claudei Torre" cogido en 
el tubo de la planta de M. Claude en Matanzas. Rev. Colegio 
de La Salle, Havana, p. 231-232. 
Tortonese, Enrico 

1958. Elenco dei Leptocardi, Ciclostomi, Pesci cartilaginei ed ossei del 
mare mediterraneo. Atti Soc. Italiani Sci. Nat. Mus. Civ. Stor. 
Nat. Milano, vol. 97, fasc. 4, pp. 309-345. 



20 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 1. — Counts and measurements of Oligopus longhursti 



Counts 


HolotTOe 

USNM 
187778 


Paratvpe 
USNM 
187779 


Paratvpe 
USNM 
191732 


Paratyne 
USNM 
191732 


Paratx-pe 
USNM 
193678 


Dorsal 

Anal 

Pectoral 

Vertebrae 

Caudal 

Vertical scale rows (approximately) 


88 
58 
25 
45 
8 
120 


89 
61 
24 
44 
8 


83 

58 
25 
45 
8 
120 


85 
61 
25 
45 
8 
120 


86 
59 
25 
44 
8 
120 


Measurements 


mm. %SL 


mm. %SL 


mm. %SL 


mm. %SL 


mm. %SL 


Standard length 

Body depth at dorsal origin 

Predorsal 

Preanal 

Head length 

Snout 

Orbit 

Upper jaw 

Greatest width of maxillary 

Greatest %vidth of head 


98.0 

27. 27. 5 

30. 30. 6 

50.5 51.5 

25. 6 26. 1 
6.9 6.0 
4.5 4.6 

15.0 15.3 
5.4 5.5 
15.0 15.3 


92.6 

23.5 25.4 
29.5 31.9 
48.5 52.4 
24. 3 26. 2 
5. 7 6. 2 
3.5 3.8 
14.0 15.1 
5. 5. 4 
13.0 14.0 


105 

25. 23. 8 
31.4 29.9 

55. 1 52. 5 
26. 2 24. 9 

6. 2 5. 9 
3.5 3.3 

14.8 14.1 
4.7 4.5 

16.0 15.2 


105 

32. 5 30. 7 

58. 54. 7 

26. 9 25. 4 
5.2 4.5 
4.2 4.0 

14.5 13.7 
4.2 4.0 

16.1 15.2 


106 

33.5 31.6 

60. 4 56. 9 

28. 5 26. 7 
6.2 5.8 
4.0 3.8 

15.5 14.6 
4.2 4.0 



Table 2. — Summary of measurements on O. diagrammus /rom three geographical 
locahties (given as percent of standard length; average in parenthesis followed hv 
range) ^ 



Measurement 



Range in standard length (mm.) 

Depth at dorsal origin 

Predorsal 

Preanal 

Head length 

Snout 

Orbit 

Upper jaw 

Maxillary width 

Head width 



Galapagos-N=2i 



23.6 
29.5 
44.1 
26.6 
5.9 
5.0 
13.8 
4.35 
14.9 



91,54 

(21.4, 25.9) 

(28.6, 30.4) 

(43.9, 44.4) 

(25.5. 27.8) 

(5.4, 6.5) 

(4.4, 5.6) 

(14.7. 13.0) 

(4.3, 4.4) 

(13.4, 16.5) 



Guadalupe-N = 13 



24.5 
29.1 
47.3 
26.0 
5.2 
4.3 
13.2 
4.4 
14.2 



74. 7-184 

2 (21.3-32.1) 

(26. 3-30. 9) 

(46.3-51.3) 

(25. 1-27. 2) 

(4. 6-6. 2) 

(3.8-1.7) 
(12.5-13.8) 

(3.9-4.7) 
(12.0-17.7) 



Peninsular Baja 

Calif. -N =8 



23.1 
29.4 
46.3 
26.9 
5.7 
4.8 
13.7 
4.5 
13.3 



34.5-81.0 
(20. 9-25. 2) 
(26.3-31.9) 
(43.8-48.1) 
(25. 3-27. 8) 
(4.^-6.6) 
(4.7-6.1) 
(12.8-14.4) 
3 (4. 0-5. 0) 
(11.1-17.3) 



* M- '^ number in parenthesis is holotype, second is paratype. 
»N=5.' 



REVIEW OF OPHIDIOID FISH GENUS OLIGOPUS — COHEN 21 



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U.S. GOVERNMENT PRINTING OFFICE: 1964-724-329 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1964 Number 3493 



NORTH AMERICAN STENOMIDAE 
(LEPIDOPTERA: GELECHIOIDEA) 



By W. Donald Duckworth 



Introduction 



This is a review of the North American moths of the family Sten- 
omidae, a large family of Microlepidoptera that is especially abundant 
in Central and South America, where it is represented by hundreds 
of species of remarkable diversity in size, shape, and color. In North 
America the family is represented by 24 species in 6 genera, widely 
distributed throughout the United States and parts of Canada. 

Formerly, identification of species in the Stenomidae rested almost 
solely on coloration and, to a lesser extent, on wing venation. Higher 
categories, for the most part, were based on wing venation. In recent 
years, it has been discovered that genitalic characters in the Sten- 
omidae, as in other groups of Lepidoptera, are of value in the separa- 
tion and definition of closely related species. It seems obvious that 
the genitalia, in addition to their usefulness in identifying species, 
have a place of primary importance in characterizing genera. The 
present study also indicates that the genitalia provide valuable 
characters for classification at family level. 

Busck (1921a) recognized the value of genital characters to higher 
category concepts and transferred the genus Setiostoma Zeller from 

23 



24 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lis 

Glyphipterygidae to Stenomidae after a careful study of their struc- 
tures. He also indicated that on the basis of the genitalia, the Sten- 
omidae and Xylorcytidae should be recognized as separate families. 

The failure of other workers to accept the evidence available from 
studies of the genitalia led to controversy and confusion. Meyrick 
admitted the value of the genitalia as an aid to classification but 
based his higher categories primarily on wing venation and other 
characters. 

Although the author is convinced that the genitalia are of primary 
importance in the problems of delimiting higher categories in the 
Stenomidae, an attempt to solve these problems without first making 
a detailed study of the Central and South American species would be 
unrealistic. A generic classification of stenomids is being undertaken 
presently as a separate study. 

The following classification uses structures of both male and female 
genitalia, as well as color and other adult characteristics. Illustra- 
tions of the genitalia of both sexes are given for each species. The 
terminology used for structures of the male and female genitalia of 
Lepidoptera is very extensive and complex. Despite this, there is a 
constant application of new names, as well as misapplication of old 
ones, by taxonomists who need names for many structures with doubt- 
ful homologies. Some workers apply the same name to structures 
similar in appearance, even in widely separated families, while others 
hesitate to apply the same name to two structures unless these have 
been shown to be strictly homologous. The majority of workers do 
not wish to contribute to further instability of nomenclature but are 
unable to await the results of morphological studies dealing with the 
homologies. The nomenclature of the genitalia in this paper follows 
the recommendations made by EHots in Tuxen's publication (1956) 
on insect genitalia. 

Biological studies and larval host plant records for North American 
stenomids are few; known records for each species are listed in the 
text. The larvae of the North American species feed chiefly on 
trees and shrubs such as oak, maple, and laurel, although one species, 
Stenoma mistrella Busck, has been found on timothy (Phleum pratense 
L.). Exotic species are known to feed on avocado, custard apple 
(Annona sp.), coffee, and cacao. 

The literature on the North American species of Stenomidae is 
somewhat confused because of the uncertainty of many authors as 
to the relationship of Stenomidae to other closely related families. 
Much of this uncertainty has come from Edward Meyrick's work 
(1880, 1889, 1890, 1913, 1915, 1922, 1925, 1928, 1929) in the Ste- 
nomidae and related groups. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 25 

As mentioned earlier, Me3'^rick's system of classification was based 
primarily on wing venation. He also tried to employ a hypothesis, 
based on a mathematical formula, to provide a convenient artificial 
system to which he could fit his classification. The desired standard 
in Meyrick's classification was an average of about 10 species to the 
genus and 50 genera to the family in the world fauna. His failure 
to accept classification based on other valuable structures, such as 
the genitalia, left many unanswered questions concerning relation- 
ships of categories within the Stenomidae and relationships of this 
family to other families. 

The relationship between Stenomidae and Xyloryctidae has pre- 
sented a difficult problem; Busck (1921a) transferred the genus 
Setiostoma ZeUer from Glyphipterygidae to Stenomidae. Forbes 
(1923) evidently felt the two families were not distinct because he 
listed them as subfamihes under the family Xyloryctidae. Clarke 
(1955a) recognized them as separate famihes: the Xyloryctidae being 
confined principally to the Old World and the Stenomidae to the 
New World. 

The genus Stenoma Zeller, upon which the family is based, is still 
a doubtful entity. The type species, Stenoma litura Zeller, is based 
on a single female now in the British Museum that, according to 
Clarke (1955a), is unlike any of the approximately 600 species that 
have been placed as congeners. Clarke (1955a) also states that this 
gi'eat mass of species is still a heterogeneity that will require close 
study, with the possible delimitation of many new genera based on 
diagnostic refinements. 

Some genera that have been considered to be doubtful with respect 
to family affinity are herein treated as Stenomidae on the basis of 
the genitaha, pending a more comprehensive study of generic rela- 
tionships throughout the family. In this paper, the family Sten- 
omidae in North America includes the genera Stenoma, Antaeotricha, 
Setiostoma, Mothonica, Menestomorpha, and Menesta. The genus 
Menesta was moved by Forbes (1923) from the family Gelechiidae to 
the Stenomidae on the basis of the genitalia, as well as other adult 
and pupal characters. 

The author gratefully acknowledges the aid of Dr. David A. Young 
of North Carolina State College, who encouraged the undertaking of 
a taxonomic problem in the Microlepidoptera and made suggestions 
on the preparation of this paper. Sincere thanks are also due Dr. 
J. F. Gates Clarke, Head Curator of Entomology, U.S. National 
Museum, for suggesting this problem, for technical advice, and for 
his kind encouragement. The loan of material from their respective 
private collections by Mr. C. P. Kimball and by Dr. Ronald Hodges 
also is gratefully acknowledged. The author also wishes to thank 



26 PROCEEDESTGS OF THE NATIONAL MUSEUM vol. ug 

Mr. Andre Pizziui for the distribution maps and Mr. Jack Scott for 
the photographic work. 

Family Stenomidae Meyrick 

Stenomidae Meyrick (in part), 1906, Trans. Roy. Soc. South Australia, vol. 30, 
p. 50; 1909, Trans. Ent. Soc. London, p. 28; 1912, Trans. Ent. Soc. Loudon, 
p. 706; 1931, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 36, p. 378.— 
Walsingham, 1912, Lepidoptera-Heterocera, vol. 4 (vol. 42 in Godman and 
Salvin, Biologia Centrali- Americana), pp. 153-187; 1913, Lepidoptera- 
Heterocera, vol. 4 (vol. 42 in Godman and Salvin, Biologia Centrali-Ameri- 
cana), pp. 188-190. — Barnes and Busck, 1920, Contrib. Nat. Hist. Lepidop. 
North America, vol. 4, p. 236. 

Stenomatidae Walsingham, 1907, Proc. U.S. Nat. Mus., voL 33, p. 214. 

Cryptolechiidae Meyrick (in part), 1883, Trans. Ent. Soc. London, p. 124. 

Xyloryctidae Meyrick (in part), 1925-1934, Exotic Microlepidoptera, vols. 1-4. 
— Forbes, 1923, Cornell Agric. Exp. Sta. Mem., vol. 68, p. 250. 

Cryptophasidae Fletcher (in part), 1929, Mem. Dept. Agric. India, Ent. Ser., 
vol. 11, pp. 1-244. 

Stenomides Meyrick, 1930, Ann. Naturhist. Mus. Wien, vol. 44, p. 233. 

Stenominae Janse, 1932, The moths of South Africa, vol. 1, p. 61. 

Type genus. — Stenoma Zeller. 

Male antenna heavily ciliated ventrally, female with shght or no 
cihation, basal segment without pecten; labial palpus compressed, 
curved gradually dorsad and extending above crown of head. Fore- 
wing with 12 veins (or 11 by fusion of veins 2 and 3) ; vein lb furcate 
at base; vein 7 separate to termen, apex or costa; veins 2-3 and 4 
separate, connate, or stalked; veins 2 and 3 fused, stalked, or separate. 
Fore wing typically much more than twice as long as wide; shorter in 
Menesta and Setiostoma. Hindwing with veins 6 and 7 stalked ; 3 and 
4 connate, stalked, or fused; vein 5 close to, connate, or fused with 4; 
typically very broad with rounded termen. 

Male genitalia: Symmetrical; harpes simple or divided into lobes, 
bearing bifm'cate setae that may be long and straight, long and re- 
curved, or short; socii and transtHla absent; aneUus a simple plate or 
with moderately or weU-developed lateral processes. Vinculum band- 
Uke, complete or incomplete. Gnathos present; uncus present, well 
developed or reduced. 

Female genitalia: Genital plate variously sclerotized or membra- 
nous; corpus bursae mth or without signum. 

Key to Species of North American Stenomidae Based on Genitalia 

1. Male 2 

Female 24 

2. Harpes with thumblike projection on costa bearing long, recurved, bifurcate 

setae (Genus Antaeotricha Zeller) 3 

Harpes without such a projection 17 



NORTH AlVIERICAN STENOMIDAE — DUCKWORTH 27 

3. Anellus with lateral lobes 4 

Anellus without lateral lobes 15 

4. Anellus with two lateral lobes 5 

Anellus \\ith four lateral lobes 11 

5. Tip of gnathos broad, notched 6 

Tip of gnathos narrow, unnotched 7 

6. Left lobe of anellus forked 1. Antaeotricha schlaegeri (Zeller) 

Left lobe of anellus simple . . 2. Antaeotricha lindseyi (Barnes and Busck) 

7. Uncus simple; lobes of anellus small, bearing setae at tips; cornuti a long mass 

of small spines 8. Antaeotricha irene (Barnes and Busck ) 

Uncus not simple; combination of characters not as above 8 

8. Aedeagus with cornuti; combination of characters not as below .... 9 
Aedeagus without cornuti; uncus with dilated tip; lateral lobes of anellus 

upright, sharply pointed . . .11. Antaeotricha haesitans (Walsingham) 

9. Vinculum produced into dorsally projecting process in front. 

13. Antaeotricha fuscorectangiilata, new species 
Vinculum not produced into dorsally projecting process 10 

10. Uncus with broadened tip; aedeagus flaring apically. 

4. Antaeotricha leiicillana (Zeller) 

LTncus with tip terminating in two short spines; aedeagus short, broad, 

truncate at tip 15. Antaeotricha nianzanitae Keifer 

11. Uncus distinctly notched at tip, aedeagus with cornuti a large irregular 

group of very small spines; with lateral recurved process. 

7. Antaeotricha furcata (Walsingham) 

Uncus without notched tip, aedeagus without cornuti and without lateral 

recurved process 12 

12. Gnathos divided into two lobes at tip 13 

Gnathos not divided into two lobes at tip 14 

13. Lobes on tip of gnathos blunt . . 3. Antaeotricha unipunctella (Clemens) 
Lobes on tip of gnathos acuminate. 

14. Antaeotricha vestalis (Barnes and Busck) 

14. Lobes of anellus unequal, dorsal lobe large, laterally curved, ^ith dense, 

brushlike group of spines on inner margin of apical third; ventral lobes 
reduced, setiferous; aedeagus without spine. 

6. Antaeotricha decorosella (Busck) 
Lobes of anellus approximately equal in size, dorsal ones with heavy spines at 
tip, ventral ones setiferous; aedeagus with apical spine. 

5. Antaeotricha osseella (Walsingham) 

15. Anellus V-shaped, uncus dilated at tip, not cleft. 

9. Antaeotricha humilis (Zeller) 
Anellus not V-shaped, uncus not dilated at tip 16 

16. Uncus attenuate, cleft at tip. 

11. Antaeotricha thoniasi (Barnes and Busck) 
Uncus truncate, not cleft at tip . . 10. Antaeotricha agrioschista (Meyrick) 

17. Harpes simple with palmate, multilobed setae on outer part of costa. 

(Genus Setiostoma Zeller) 18 

Harpes simple or compound without palmate, multilobed setae on outer part 

of costa 19 

18. Cornuti consisting of more than one large single spine. 

2. Setiostoma ferualdella Riley 
Cornuti consisting of one large single spine. 

1. Setiostoma xanthobasis Zeller 



28 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

19. Harpes compound 20 

Harpes simple 21 

20. Sacculus with prominent bifurcate lobe. 

(Genus Mothonica Walsingham) 1. Mothonica kimballi, new species 
Sacculus a large JBeshy lobe . . (Genus Menestomorpha Walsingham) 

1. Menestomorpha oblongata (Walsingham) 

21. Uncus reduced (Genus Stenoma Zeller) 22 

Uncus large, tapering, curved ventrad, lying parallel to gnathos. 

(Genus Menesta Clemens) 23 

22. Aedeagus large, flared apically; cornuti two clusters of very heavy spines. 

1. Stenoma mistrella Busck 
Aedeagus small, broad; cornuti one small cluster of heavy spines. 

2. Stenoma crambitella Walsingham 

23. Cornuti consisting of two heavy spines. 

1. Menesta tortriciformella Clemens 
Cornuti consisting of one heavy spine. . . 2. Menesta melanella Murtfeldt 

24. Anterior apophyses fused to genital plate or very reduced. 

(Genus Aniaeotricha Zeller) 25 
Anterior apophyses free, well developed 37 

25. Corpus bursae with signum 28 

Corpus bursae without signum 26 

26. Corpus bursae corrugated, genital plate bandlike. 

15. Antaeotricha manzanitae Keifer 
Corpus bursae not corrugated, genital plate not bandlike 27 

27. Genital plate reduced, lamella antevaginalis and postvaginalis not differenti- 

ated, ductus bursae short, sclerotized, corpus bursae small. 

4. Antaeotricha leucillana (Zeller) 

Genital plate well developed, lamella antevaginalis small, forming triangular 

flap over ostium 12. Antaeotricha haesitans (Walsingham) 

28. Genital plate reduced to simple band, signum with single large spine . . 29 
Genital plate more complex, signum a small- to large-toothed plate . . 32 

29. Anterior apophyses fused to genital plate. 

11. Antaeotricha thomasi (Barnes and Busck) 
Anterior apophyses small, not fused to genital plate 30 

30. Ostium bursae with a pouchlike evagination between ostium and inception 

of ductus seminalis . . 13. Antaeotricha fuscorectangulata, new species 
Ostium bursae without such an opening 31 

31. Genital plate excavated along posterior margin, ductus bursae short. 

10. Antaeotricha agrioschista (Meyrick) 
Genital plate not excavated, ductus bursae long. 

9. Antaeotricha humilis (Zeller) 

32. Ostium opening near center of genital plate 33 

Ostium opening near anterior margin of genital plate 36 

33. Ductus bursae sclerotized from midpoint to corpus bursae 

8. Antaeotricha irene (Barnes and Busck) 
Ductus bursae not sclerotized 34 

34. Posterior margin of genital plate cleft medially. 

5. Antaeotricha osseella (Walsingham) 
Posterior margin of genital plate not cleft 35 

35. Genital plate with median T-shaped process extending over ostium. 

3. Antaeotricha unipunctella (Clemens) 
Genital plate without process 14. Antaeotricha vestalis (Zeller) 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 29 

36. Genital plate with large, median, elevated area directly posterior to os- 

tium 37 

Genital plate without such an area . . 6. Antaeotrieha decorosella (Busck) 

37. Ductus bursae thick, short, approximately same length as corpus bursae. 

7. Antaeotricha furcata (Walsingham) 
Ductus bursae slender, long, approximately twice length of corpus bursae. 

1. Antaeotricha schlaegeri (Zeller) 
or 2. Antaeotricha lindseyi (Barnesand Busck) 

38. Genital plate membraneous (Genus Setiostoma Zeller) 39 

Genital plate not membraneous 40 

39. Corpus bursae with signum 1. Setiostoma xanthobasis Zeller 

Corpus bursae without signum 2. Setiostoma fcrnaldella Riley 

40. Genital plate fused to eighth sternum (Genus Sienoma Zeller) 41 

Genital plate not fused to eighth sternum 42 

41. Corpus bursae with one lightly sclerotized signum. 

1. Stenonia mistrella Busck 
Corpus bursae with two lightly sclerotized, dentate signa. 

2. Stenoma crambitella Walsingham 

42. Signum large, dumbbell-shaped, dentate. 

1. INIothonica kimballi, new species 
Signum not as above 43 

43. Genital plate reduced to small, median, triangular sclerite. 

(Genus Menesta Clemens) 44 
Genital plate reduced to simple liplike band surrounding ostium. 

1. Menestomorpha oblongata Walsingham 

44. Corpus bursae with heavily sclerotized, cross-shaped signum with median 

projecting lobe 1. Menesta tortriciformella Clemens 

Corpus bursae with heavily sclerotized, rectangular signum with median 
projecting lobe 2. Menesta melanella Murtfeldt 

Genus Antaeotricha Zeller 

Antaeotricha Zeller, 1854, Linn. Entom., vol. 9, p. 390. 

1. Antaeotricha schlaegeri (Zeller) 

Figures 1, 24; Plate la; Map 1 

Cryptolechia schlaegeri Zeller, 1854, Linn. Entom., vol. 9, p. 372; 1855, Linn. 
Entom., vol. 10, p. 158; 1873, Verh. Zool.-Bot. Ges. Wien, vol. 23, p. 246. 

Stenoma schlaegeri Walsingham, 1889, Insect Life, vol. 2, p. 152. — Barnes and 
Busck, 1920, Contrib. Nat. Hist. Lep. North America, vol. 4, p. 238. — 
Forbes, 1923, Cornell Agric. Exp. Sta. Mem., vol. 68, p. 252. 

Antaeotricha schlaegeri Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 28. 

Alar expanse 21-27 mm. 

Color: Face white; palpus white, sprinkled with dusky scales. 
Legs white; foreleg smoky; midleg with dark tarsi ringed with light 
scales. Thorax white dorsally with brownish-black tuft posteriorly. 
Abdomen white. Forewings white, basal half with conspicuous, 
mixed light and dark brown, somewhat raised patch of scales on 
hindmargin from basal angle almost to middle of wing, apical 
half with irregular discal area of dusky scales; postmedial and 



30 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

subterminal bands broad, even; duslrjr terminal band very narrow, 
cut into spots; black line, usually broken, in base of cilia. Hindwing 
very pale tan with white cilia. 

Male genitalia: Uncus simple, curved ventrad; gnathos very broad, 
broadly notched at tip; harpes each with costa bearing a thumblike 
projection with long, recurved, bifurcate setae; -vdnculum complete; 
anellus with two large, upright lateral lobes, one forked, the other 
simple; aedeagus with apex acute, cornuti absent. 

Female genitalia: Genital plate large, lameUa antevaginalis a small, 
semicircular plate; lamella postvaginalis larger, with median elevated 
area; ostium bursae and ductus bursae membranous; corpus bursae 
with signum large, dentate; inception of ductus seminalis near ostium. 
Anterior apophyses fused to genital plate. 

Type: In the British Museum (Natural History). 

Type locality: New York. 

Food plant: Quercus alba L. 

Distribution: Quebec: Meach Lake (May, June), new york: 
Allegany State Park (June, July, Aug.) ; Flatbush (June) ; Ilion (June). 
MASSACHUSETTS: Martha's Vineyard (June); Vineyard Haven (June). 
PENNSYLVANIA: Bcavcr Co. (May); New Brighton (May, June); 
Pittsburgh (May), new jersey: Essex Co. Park (May, June). 
DELAWARE: Ncw Castlc (June). Maryland: Plimamers Island (May); 
Riverdale (June), district of Columbia: Washington (Jime). 
Virginia: Falls Church (May, Aug.); Grange Camp (May), north 
Carolina: Raleigh (April, June, Aug.). Arkansas: Oracle (July). 
MISSOURI: St. Louis (June). Illinois: Quincy (May) ; Decatur (May). 
10 wa: Iowa City (June); Sioux City (May); Homestead (May). 
TEXAS: Waco (May); Dallas. Arizona: Huachuca Mts. (Oct.); 
Palmerlee; Oracle (July). 

This species is very similar to Antaeotricha lindseyi. Possibly the 
two are conspecific, but until series of A. lindseyi can be obtained and 
the variation studied, I prefer to recognize them as distinct species. 
They may be distinguised by differences in the shapes of the aedeagus 
and of one of the anellar lobes. Significant differences in the female 
genitaUa have not been found. 

2. Antaeotricha lindseyi (Barnes and Busck) 

Figure 2; Plate lb; Map 1 
Stenoma lindseyi Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North America, 

vol. 4, p. 239. 
Antaeotricha lindseyi Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 25. 

Alar expanse 25-28 mm. 

Color: Face, palpus, and legs white. Thorax white dorsally with 
brown tuft of scales posteriorly. Abdomen white. Forewing longer, 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 31 

narrower, and more pointed than in A. schlaegeri, and with dark area 
on hindmargin (which is interrupted near middle of forewing in A. 
schlaegeri) continued to anal angle. Hindwing of male dark brownish, 
considerably darker than those of A. schlaegeri. 

Male genitalia: As in A. schlaegeri, except anellar lobe hooked in- 
stead of forked, and aedeagus shaped differently. 

Female genitalia: As in A. schlaegeri. 

Type: In the United States National Museum. 

Type locality: Paradise, Cochise County, Arizona. 

Food plant: Unknown. 

Distribution: Arizona: Prescott (Sept.); Paradise, Cochise Co. 
(July, Sept.); Gila Co. (June); Palmerlee; Yavapai Co. (Sept.); 
(Huachuca Mts.; Baboquivarii Mts. (July); Mohave Co. (Sept.); 
Dewey; Redington. new Mexico: Rincon (June); Ft. Wingate 
(June) . 

This species is closely related to A. schlaegeri; the distinguishing 
characters have been noted under that species. 

3. Antaeotricha unipunctella (Clemens) 

Figures 3, 25; Plate Ic; Map 2 
Brachiloma unipunctella Clemens, 1863, Proc. Ent. Soc. Philadelphia, vol. 2, p. 

126; 1872, The Tineina of North America, p. 232.— Busck, 1903a, Proc. 

Ent. Soc. Washington, vol. 5, p. 214. 
Cryptolechia lithosina Zeller, 1873, Verh. Zool.-Bot. Ges. Wien, vol. 23, p. 244. 
Harpalyce iortricella Chambers, 1874, Canadian Ent., vol. 6, p. 235. 
Ide Iortricella Chambers, 1877, Bull. U.S. Geol. Surv., vol. 3, p. 122, 141. 
Ide lithosina Walsingham, 1889, Insect Life, vol. 2, p. 155. 
Slenoma unipunctella Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North 

America, vol. 4, p. 238. — Forbes, 1923, Cornell Agric. Exp. Sta. Mem., vol. 

68, p. 253. 
Antaeotricha unipunctella Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 30. 

Alar expanse 20-22 mm. 

Color: Face and palpus white. Foreleg bro"\vnish; mid- and hindleg 
white. Thorax stramineous dorsally. Forewing stramineous; discal 
cell with or without one or two brownish dots apically. Hindwing 
white, 

Male genitalia: Uncus a long, narrow stem beyond dilated base, 
arched and spatulate at apex. Gnathos broad, divided into two 
rounded lobes at tip; harpes as m A. schlaegeri; vinculum complete, 
arching in front; anellus mth four lateral lobes, two on each side of 
aedeagus, hindlobes longer and with heavy spines apically; aedeagus 
dilated at base, tapering to smoothly rounded tip, cornuti absent. 

Female genitalia: Genital plate wide, short; ostimn bursae sclero- 
tized; ductus bursae membranous; corpus bursae with signum large, 
dentate; inception of ductus seminalis near ostium. 

Type: In Academy of Natural Sciences of Philadelphia. 

726-597—64 2 



32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

Type locality: Pennsylvania {B. unipunctella) , Texas (C. lithosina, 
H. tortricella) . 

Food plant: Quercus sp. In the USNM collection there is one 
specimen that has been reared from chestnut. 

Distribution: Florida: Fort Myers (Apdl); Cocoanut Grove; St. 
Petersburg (Oct.) ; Enterprise (May) ; Hastings (Sept.) ; Oneco (May) ; 
Weeki Wachee Springs (May); Sarasota (May). Louisiana: Natchi- 
toches Parish (Aug.). texas: Kerrville (April, May, June, Aug.); 
Burnett Co. (Sept., Oct.); Belfrage; Shovel Mt. (July). Arizona: 
Yavapai Co.; Ft. Grant (July). 

This species is very similar to A. decorosella in both coloration and 
genitalia; however, the fore wings are stramineous in color, the male 
genitalia have a much larger and more broadly notched gnathos, the 
aedeagus is shaped differently, and the female genital opening is in 
the center of the genital plate rather than on the anterior edge. 

4. Antaeotricha leucillana (Zeller) 

Figures 4, 26; Plate Id; Map 3 

Cryptolechia leucillana Zeller, 1854, Linn. Entom., vol. 9, p. 370. 

Cryptolechia algidella Walker, 1864, List of the specimens of lepidopterous insects 

in the collection of the British Museum, vol. 29, p. 710. (New synonymy.) 
Stenoma leucillana Walsingham, 1889, Insect Life, vol. 2, p. 153. 
Stenoma algidella Walsingham, 1889, Insect Life, vol. 2, p. 153. — Barnes and 

Busck, 1920, Contrib. Nat. Hist. Lep. North America, vol. 4, p. 238. 
Antaeotricha leucillana Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 25. 

Alar expanse 15-23 mm. 

Color: Face and palpus white, sprinkled with brown. Legs white; 
forelegs covered with dark scales. Thorax white dorsally with 
brownish black tuft posteriorly as in A. schlaegeri but former with 
brown cilia on apical half. 

Male genitalia: Uncus with broadened, bifid tip; gnathos simple, 
slender, ending in a kind of beak; harpes as in A. schlaegeri, vinculum 
complete, anellus with two rounded, lateral lobes; aedeagus large, 
flaring apically with ventral half extending farther than dorsal and 
forming a broad lip; cornuti a cluster of many medium-sized spines. 

Female genitalia: Genital plate reduced to simple band; ostium 
bursae and ductus bursae short, sclerotized; corpus bursae small; 
signum absent; inception of ductus seminalis near ostium. 

Type: Unknown. 

Type locality: Georgia (C. leucillana), Nova Scotia (C. algidella). 

Food plants: Pyracantha crenulata (Roxb.), Malus sp., Vaccinium 
corymbosum L., Acer sp. Frost (1931) gives an account of the Hfe 
history of S. algidella (= A. leucillana). 

Distribution: new Hampshire: Hampton (June, July); White 
Mts. MASSACHUSETTS: Martha's Vineyard (July), new york: 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 33 

Orient (Aug., Sept., Oct.); McLean Bogs Reserve (May, June); 
Ithaca (May, June, Aug.) ; Boreas River, Essex Co. (July) ; Mattituck 
(June) ; Trenton Falls (June) ; Rock City (June) ; West Falls (Aug.) ; 
Buffalo (July); E. Aui-ora (May, June); Horseheads (May); Monroe 
Co. (June). PENNSYLVANIA: South Gibson (Aug.); Oak Station 
(May, July); New Brighton (April, May, June); Pittsburgh (May, 
Aug.); Beaver Co. (Alay). new jersey: Reach (Aug.); Anglesea 
(May) ; Park (Aug.) ; Whitesbog (May, Oct.) ; Oaldand (Aug.) ; New 
Lisbon (July). Maryland: Plummers Island (June, Aug.); Hyatts- 
ville (May); Cabin John (June), district of Columbia: Washington 
(May, June). Virginia: Shenandoah (Aug.); Stafford (May). 
WEST VIRGINIA: White Sulphur Springs (July), north Carolina: 
Tryon (May); Southern Pines (July, Aug.); White Lake (July). 
SOUTH Carolina: Oconee (Aug.). Florida: Gainesville (July); 
St. Petersburg (Jan.); Vero Beach (Feb., March, Oct., Nov.); Oneco 
(June); Weeki Wachee Springs (June); Siesta Key (May, June); 
Bradenton (Aug.); Pensacola (March). Georgia: Savannah (July). 
ALABAMA: Camp Rucker (April); Flatwood (June); Leroy (June). 
Mississippi: Biloxi (June); Ocean Springs (April); Bolton (Aug.). 
ARKANSAS: Sulphur City (Aug.). Missouri: Mountain Grove 
(July). KANSAS: Lawrence (April, May). Illinois: Decatur 
(March, April, Maj'-). iowa: Iowa City (JuljO; Sioux City (June, 
July); Homestead (May), ohio: Cuyahoga Co. (May); Dayton 
(May). Manitoba: Cartwright. texas: Denver (June); Browns- 
ville (June); San Benito (Aug.); Kerrville (June); Tiger Mill; Victoria 
(Sept.); Shovel Mt. (July); Mathis (Aug.); Laredo (Sept.); Mercedes; 
Dallas (April). Oregon: Ritter, Grant Co. (June). Louisiana: 
Sam Houston State Park (Aug). 

This species is very similar to A. schlaegeri in coloration but it 
is generally smaller. The presence of an uncus with a broadened 
bifid tip and a broad, apically flared aedeagus in the male genitalia, 
and a reduced genital plate and small corpus bursae in the female 
genitalia readily separate this species from A. schlaegeri. 

The name of this species has been in doubt for a number of years. 
The type locality for Walker's C. algidella is Nova Scotia and for 
ZeUer's C. leiwillana is Georgia. There is, according to Walsingham 
(1889), some color variation between the southern and northern 
limits of distribution and this variation is responsible for Walker's 
description of C. algidella. Although the location of Zeller's type is 
uncertain, a Zeller specimen from Texas labeled C. leucillana in the 
British Museum is identical to Walker's type of C. algidella. Ob^dously 
this problem cannot be resolved adequately until Zeller's type of 
C. leucillana is studied and compared with that of Walker's C. algi- 
della; however, since indirect evidence indicates that both names 



34 PROCEEDENTGS OF THE NATIONAL MUSEUM vol. lie 

apply to the same species, and, since C. leucillana is the older name, 
the latter is adopted provisionally here. 

5. Antaeotricha osseella (Walsingham) 

Figures 5, 27; Plate le; Map 4 

Ide osseella Walsingham, 1889, Insect Life, vol. 2, p. 155. 

Brachyloma querciella Busck, 1908, Proc. Ent. Soc. Washington, vol 10, p. 111. 

(New synonymy). 
Stenoma querciella Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North 

America, vol. 4, p. 238. — Forbes, 1923, Cornell Agric. Exp. Sta. Mem., vol. 

68, p. 253.— Busck, 1925, Proc. Ent. Soc. Washington, vol. 27, p. 48. 
Stenoma osseella Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North America, 

vol. 4, p. 239. 
Antaeotricha osseella Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 26. 
Antaeotricha querciella Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 27. 

Alar expanse 22-24 mm. 

Color: Face and palpus whitish sprinkled with brown scales. Legs 
whitish, forelegs dark fuscous anteriorly. Thorax brown dorsaliy. 
Abdomen ochreous. Forewing brown with strong lustre; with two 
blacldsh dots at end of cell, anterior one more proximal than posterior 
one. Hindwing light tan with white cUia. 

Male genitalia: Uncus similar to A. unipunctella, with spatulate 
tip, but not as arched. Gnathos broadly notched at tip; harpes as 
in A. schlaegeri; vinculum complete, arching in front ; anellus as in A. 
unipunctella but larger and notched medially; aedeagus tapering to 
rounded apex bearing a small apical spine. Cornuti absent. 

Female genitalia: Genital plate wide, short, mth median slot. 
Ostium bursae large; ductus bursae long, membranous; corpus 
bm-sae narrow with signum large, dentate; inception of ductus semi- 
nalis near ostium. 

Type: In the British Museum (Natural History). 

Type locality: California (7. osseella), Montclair, New Jersey 
(B. querciella). 

Food plant: Quercus alba L., Quercus muehlenbergii Engelm. 

Distribution: new york: Ithaca (July, Aug., Sept.). new jersey: 
Lakehurst (Aug., Sept.); Bro\vn's Mills (July), north Carolina: 
Highlands (June, Aug.); Brevard (Aug.). south Carolina: Oconee 
(Aug., Sept.); GreenvUle; Anderson (June), west Virginia: White 
Sulphur Springs (July). Maryland: Hyattsville (May, June), dis- 
trict OF COLUMBIA: Washington (Sept.). Massachusetts: Newton 
Highlands. Pennsylvania: New Brighton (July, Aug.). Illinois: 
Decatur (July). Arkansas: Washington Co. (July, Aug.). texas: 
Burnet Co. Missouri: Columbia. 

This species is similar in size to A. decorosella and A. unipunctella, 
but the coloration is distinct. The spine on the tip of the aedeagus 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 35 

in the male and the median slot in the genital plate in the female are 
distinctive genital characters. 

The identity of this species has been in doubt for some time. 
Keifer (1937) discusses the status of the name osseella and states: 
"At the present time I know of no further published elucidation of 
the application of the name osseella." I have examined the genitalia 
of Busck's type of B. querciella and photographs of the genitalia of 
Walsingham's type of /. osseella and found the two identical. Since 
Walsingham's is the older name, Busck's B. querciella must fall. 

6. Antaeotricha decorosella (Busck) 

Figures 6, 30; Plate If; Map 4 

Brachyloma decorosella Busck, 1908, Proc. Ent. Soc. Washington, vol. 10, p. 111. 
Stenoma decorosella [sic] Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North 

America, vol. 4, p. 256. 
Stenoma decorella [sic] Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North 

America, vol. 4, p. 256. 
Antaeotricha decorosella Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 21. 

Alar expanse 22-24 mm. 

Color: Face whitish ochreous; palpus brownish ochreous. Foreleg 
rich brown; midleg a somewhat lighter shade; hindleg white. Thorax 
dark brown dorsally. Abdomen whitish ochreous. Forewing dark 
brown; costal edge narrowly light ochreous; cell with barely percepti- 
ble darker brown apical spot. Hindmng whitish fuscous with cilia 
ochreous. 

Male genitalia: Uncus curved, dilated at tip and only slightly in- 
dented; gnathos well developed and notched at tip; harpes as in A. 
schlaegeri; anellus with four lobes, ventral ones small amd thumblike, 
bearing several setae at their apex, dorsal ones large, laterally curved, 
with dense, brushlike group of spines on inner inargin of apical third. 
Aedeagus long, with slightly pointed apex; cornuti absent. 

Female genitaha: Genital plate large, lamella antevaginaUs a small 
liplike band, lamella postvaginalis much larger, slightly elevated in 
center; ostium bursae large, slightly sclerotized; ductus bursae long, 
membranous; corpus bursae with large, toothed signum; inception of 
ductus seminalis near ostium. 

Type: In the United States National Museum. 

Type locaUty: Montclair, New Jersey. 

Food plant: Quercus ilicifolia Wang, Ouercus marilandica Muench. 

Distribution: north Carolina: Tryon (Aug.). new jersey: New 
Lisbon (Aug.) ; Lakehurst (July, Aug.) . Florida : St. Petersburg 
(April). MASSACHUSETTS: Martha's Vineyard (Aug.). 

This species is similar to A. unipunctella, but it differs in the presence 
of a dense, brushlike group of spines on the dorsal lobes of the anellus 



36 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

in the male genitalia and in the location of the genital opening at the 
anterior edge of the genital plate in the female. 

7. Antaeotricha furcata (Walsingham) 

Figures 7, 28; Plate 2a; Map 4 
Stenoma furcata Walsingham, 1889, Insect Life, vol. 2, p. 153. — Barnes and 

Busck, 1920, Contrib. Nat. Hist. Lep. North America, vol. 4, p. 238. 
Antaeotricha furcata Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 23. 

Alar expanse 27-30 mm. 

Color: Face and palpus white. Legs whitish, unspotted. Thorax 
whitish slightly sprinkled with dusky scales dorsally, without tuft of 
dark scales posteriorly. Abdomen cinereous. Forewing white; 
patch of dark, raised scales at base of posterior edge as in A. schlaegeri; 
with brownish-grey scales extending from this to anal angle behind 
discal cell; with faint greyish clouds and spots at apex of cell and 
brownish-grey transverse line between this and apical margin. Hind- 
wing dark cinereous in male; pale greyish ochreous in female. Cilia 
white, shaded with grey at tip ; few divided black scales present. 

Male genitalia: Uncus curved, widened, and distinctly notched at 
tip; gnathos somewhat reduced in size, notched at tip; harpes as in 
A. schlaegeri; anellus with four lobes, ventral ones small with pointed 
apex bearing several setae, dorsal ones larger, truncate, bearing 
several setae at apex; aedeagus long, a long recurved process near apex, 
apex pointed; cornuti a large cluster of very small spines. 

Female genitalia: Genital plate large, lamella antevaginalis 
moderately large and liplike, lamella postvaginalis with two lateral de- 
pressions and median elevated area; ostium bursae large; ductus bur- 
sae short, membranous ; corpus bursae with toothed signum ; inception 
of ductus seminalis near ostium. 

Type: In the British Museum (Natural History). 

Type locality: Arizona. 

Food plant: Unknown. 

Distribution: Arizona: Paradise, Cochise Co. (May, June); Ma- 
dera Canyon, Santa Rita Mts. (Aug., Sept., Oct.); Morrison; Nogales 
(May). NEW MEXICO: Ruidosa Canyon (July). 

This species is similar to A. schlaegeri and A. lindseyi in color, but 
the distinctive aedeagus and notched uncus serve to separate the 
males, while the characteristic genital plate and the short, thick ductus 
bursae separate the females. 

8. Antaeotricha Irene (Barnes and Busck) 

Figures 8, 8a, 29; Plate 2b; Map 5 
Stenoma irene Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North America, 

vol. 4, p. 239. 
Antaeotricha irene Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 24. 

Alar expanse 19-20 mm. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 37 

Color: Face white; palpus white sprinkled with brown exteriorly. 
Legs white; foreleg with dark scales on outer side, mid- and hindlegs 
with dusky tarsi. Thorax ranging from dark fuscous to white dorsally 
with somewhat large, dark, rounded tuft posteriorly. Abdomen 
white sprinkled dorsally with dusky scales. Females with forewing 
white, extreme base of posterior edge fuscous; conspicuous dark spot 
at apical two-thirds; more distal, lighter, in-egular band extending to 
posterior edge; and still more distal, very faint, narrow, transverse, 
outwardly curved line across apical part of wing. In some specimens 
a cloudy spot at midlength of posterior edge. Male with entire basal 
portion of forewing blackish brown and, in addition to markings of 
female, with additional cloudy, ill-defined, more or less transverse 
areas on middle of forewing and across apical third; also an inter- 
rupted line of small black marginal dots on apex. Hindwing light 
whitish fuscous ; cilia white. 

Male genitaUa : Uncus simple ; gnathos well developed and pointed 
at tip; harpes as in A. schlaegeri; anellus narrow, with two small 
lobes; aedeagus large, sharply pointed at apex; cornuti a long mass 
of small spines. 

Female genitalia: Genital plate a small band; ostium bm-sae sclerot- 
ized, opening near center of genital plate; ductus bursae long, mem- 
branous from ostium bursae to midlength, sclerotized from midlength 
to corpus bursae; corpus bursae with dentate signum; inception of 
ductus seminalis near ostium. 

Type: In the United States National Museum. 

Type locality: Brownsville, Texas. 

Food plant: S'ic^a spp. 

Distribution: Texas: Brownsville (Jan., June^; San Benito (July, 
Aug., Sept.). 

This species is intermediate between A. leucillana and A. vesfulis 
in color, differing, in the male genitalia, by the presence of a simple 
uncus and a large cornutus in the aedeagus, and, in the female, by 
the lower part of the ductus bursae being sclerotized. 

9. Antaeotricha humilis (Zeller) 

Figures 11, 31; Plate 2c; Map 5 

Cryptolechia humilis Zeller, 1855, Linn. Entom., vol. 10, p. 156. 

Cryptolechia nebeculosa Zeller, 1873, Verb. Zool.-Bot. Ges. Wien, vol. 23, p. 245. 

Harpalyce canusella Chambers, 1874, Canadian Ent., vol. 6, p. 235. 

Sienoma humilis Walsingham, 1889, Insect Life, vol. 2, p. 154. — Barnes and 

Busck, 1920, Contrib. Nat. Hist. Lep. North America, vol. 4, p. 238. 
Antaeotricha humilis Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 24. 

Alar expanse 12-15 mm. 

Color: Face white sprinkled with brown; palpus white, sprinkled 
with brown exteriorly. Foreleg brown, tarsi ringed with white; 



38 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

midleg white sprinkled with brown; hindleg white. Thorax brown 
dorsally, without posterior tuft. Abdomen whitish ochreous. Fore- 
wing ash grey with three diffuse brown spots on anterior margin; 
three dark dots in midline, middle dot less distinct; two transverse 
bands on posterior margin; faint, curved subterminal band and row 
of dark terminal dots. Hindwing light tan; cilia tan. 

Male genitalia: Uncus curved, dilated at tip; gnathos well devel- 
oped, somewhat pointed at tip; harpes as in A. schlaegeri; anellus 
V-shaped, without lobes; aedeagus small, with slightly pointed apex; 
cornuti absent. 

Female genitalia: Genital plate small, lamella antevaginaUs and 
lamella postvaginalis not differentiated. Ostium bursae sclerotized, 
opening at posterior margin of genital plate; ductus bursae long, 
membranous; corpus bursae with large sclerotized signum; inception 
of ductus seminalis near ostium; anterior apophyses short, not fused 
to genital plate. 

Type: Not known. 

Type locality: South Carolina {C. humilis), Texas (C. nebeculosa, 
H. canuselia). 

Food plant: Quercus sp. This species has been reared by the 
author from larvae collected on oak in North Carolina in July. The 
larvae tie two leaves together and feed between them. Pupation 
occurs in the space between the two tied leaves. These observations 
are casual and a more thorough study of the life history of this species 
is planned. 

Distribution : north Carolina : Brevard (June) ; Tryon (June) ; 
Southern Pines (May, June, Sept.) ; Kinston (July) . south Carolina : 
Oconee (Aug., Sept.). Florida: Gainesville (July); Venice (Dec); 
Lake Alfred (July); Royal Palm State Park; Paradise Key (March); 
Miami; Glenwood ; Vero Beach (Feb.) ; St. Petersburg (May) ; Panacea 
(Aug.); Pensacola (Sept.). Missouri: Kirkwood (April). Missis- 
sippi: Bay St. Louis (June); Clinton (May). Tennessee: Monteagle 
(July). VIRGINIA: Cape Henry (Aug.); Falls Church (Aug.). Illi- 
nois: Oconee (July, Aug.). Maryland: Plummers Island (May); 
Hyattsville (July), texas: Waco; Kerrville (April). Indiana: Hess- 
ville (May), new jersey: Anglesea (May, June); Lakehurst (May). 
LOUISIANA : Vowell's Mill (April) . 

This species is similar to A. vestalis in size and coloration, but 
closer to A. thomasi in structures of the male and female genitalia. 
It is readily separable by the V-shaped anellus without lateral lobes 
in the male and by the presence of free anterior apophyses in the 
female. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 39 

10. Antaeotricha agrioschista (Meyrick) 

Figures 10, 10a, 34; Plate 2d; Map 5 

Stenoma agrioschista Meyrick, 1927, Exotic Microlepidoptera, vol. 3, p. 365. — 
Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 32. 

Antaeotricha agrioschista Clarke, 1955b, Catalogue of the type specimens of 
Microlepidoptera in the British Museum (Natural History) described by 
Edward Meyrick, vol. 2, p. 16, pi. 8, figs. 1-lb. 

Alar expanse 20-21 mm. 

Color: Antennae mixed brown and white. Face white, lightly- 
shaded with brown; palpus white sprinkled with brown, base of 
second segment and subapical band of apical segment brown. Legs 
white, shaded with brown, progressively lighter to rear. Thorax and 
tegula white heavily shaded with brown. Abdomen white shaded 
with brown. Forewing white overlaid brown, sprinkled with fuscous; 
short transverse fuscous line from base of costa; three oblique trans- 
verse fuscous lines from costa, one at basal fourth, one at middle and 
one at apical fourth; cUia white mixed with brown. Hindwing light 
brown; cilia whitish with brown subbasal line. 

Male genitalia: Uncus arched, area near middle expanded ventrally 
and laterally flattened; gnathos broad, rounded at tip; vinculum 
complete, arched in front, notched at apex of arch; anellus without 
lateral lobes; aedeagus slender; cornuti a long cluster of heavy spines. 

Female genitalia: Genital plate with excavation on posterior 
margin. Ostium bursae large, sclerotized; ductus bursae short, 
membranous; corpus bursae with large sclerotized signum; anterior 
apophyses short, not fused to genital plate. 

Type: In the British Museum (Natural History). 

Type locaHty: Alpine, Texas, 5000-8000 ft. 

Food plant: Unknown. 

Distribution: texas: Alpine, 5000-8000 ft. (April, May, June). 

This species is nearest A. humilis, as noted by MejTick in the 
original description. The presence of cornuti in the aedeagus and 
the notched vinculum in A. agrioschista readily separate the males, 
while the excavation on the posterior margin of the genital plate 
separates the females. 

11. Antaeotricha thomasi (Barnes and Busck) 

Figures 12, 32; Plate 2e; Map 6 

Stenoma thomasi Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North America, 

vol. 4, p. 240. 
Antaeotricha thornasi Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 29. 

Alar expanse 21-24 mm. 

Color: Face and palpus creamy white. Legs white sprinkled with 
dusky scales; tarsal joints dusky. Thorax and abdomen creamy 



40 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii« 

white. Forewing yellowish creamy white. Hindwing creamy white. 
Cilia white. 

Male genitalia: Uncus cm"ved, attenuated, and cleft at tip; gnathos 
somewhat pointed at tip; harpes as in A. schlaegeri; anellus without 
distinct lobes, aedeagus bluntly pointed at apex; cornuti a group of 
heavy spines. 

Female genitalia: Genital plate small, bandlike; lamella antevagi- 
nalis and postvaginalis similar in size and shape forming liplike struc- 
ture; ostium bursae and ductus bursae sclerotized; ductus bursae 
short; corpus bursae large, membranous, with large signum; inception 
of ductus seminalis near ostium. 

Type: In the United States National Museum. 

Type locahty: Palmerlee, Arizona. 

Food plant: Unknown. 

Distribution: Arizona: Paradise, Cochise Co. (Aug.); Huachuca 
Mts. NEW MEXICO: Albuquerque (July). Colorado: Kock Creek 
Canyon (Aug.). 

This species is similar to A. vestalis in size and color, but the at- 
tenuated cleft uncus in the male distinguishes A. thomasi from all 
other described North American species of this genus. 

12. Antaeotricha haesitans (Walsingham) 

Figures 13, 35; Plate 2f; Map 6 
Aedemoses haesitans Walsingham, 1912, Lepidoptera-Heterocera, vol. 4 (vol. 42 

in Godman and Salvin, Biologia Centrali- Americana), p. 154. 
Aedemoses hessitans [sic] Heinrich, 1921, Journ. Agric. Res., vol. 20, p. 816. 
Antaeotricha hesitans [sicj Busck, 1934, in Lepidopterorum catalogus, vol. 67, 

p. 23. 

Alar expanse 12 mm. 

Color: Face and palpus ochreous, palpus shaded with fuscous; legs 
pale ochreous. Thorax ochreous dorsally. Abdomen ochreous. 
Forewing pale ochreous with two oblique fuscous lines on apical 
half, more basal one extending from origin of costal cilia directly to 
tornus, apical one extending from dot near tornus obliquely to costa; 
minute fuscous spot at base of cell and another at its apex; a few 
minute fuscous dots at wing apex near base of cilia. Hindwing 
ochreous, cilia ochreous. 

Male genitalia: Uncus long, broadened apically, somewhat indented, 
gnathos well developed, sharply pointed at tip; harpes as in A. 
schlaegeri; vinculum complete; anellus with two upright, sharply 
pointed lobes curving laterad, one on either side of aedeagus ; aedeagus 
long, thin, with many sharp processes at apex; cornuti absent. 

Female genitalia: Genital plate short, lamella antevaginalis small, 
forming triangular flap over ostium, lamella postvaginalis larger, 
smooth; ostium bursae and ductus bursae membranous; corpus bursae 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 41 

membranous, without signum; inception of ductus seminalis near 
ostium. 

Type : In the British Museum (Natural History) . 

Type locahty: Presidio, Durango, Mexico. 

Food plant: Pitheoellobium flexicaule Benth. Heinrich (1921) 
states that the larva is a leaf-tyer, feeding on the food plant by bind- 
ing together several leaves and feeding within the tie, eating first the 
epidermis and later all but the veins of the leaves. Pupation occurs 
within the tie, the pupa bemg naked and attached to one of the 
leaves by a strand of silk. 

Distribution: texas: Brownsville (March, April, June, Aug.). 

Walsingham described a new genus, Aedemoses, and a new species 
based on a unique female with hindlegs missing, collected at Presidio, 
Durango, Mexico. 

13. Antaeotricha fuscorectangulata, new species 

Figures 14, 14a, 33; Plate 3a; Map 7 

Alar expanse 17-19 mm. 

Color: Antenna brown, irregularly shaded with white. Face whit- 
ish, shaded lightly with brown; second segment of palpus white; 
exterior brown basally; apical segment white. Legs whitish shaded 
with brown, tarsi fuscous, inner side of foreleg fuscous. Thorax 
ochreous, tegula whitish. Abdomen fuscous dorsally; ochreous 
ventrally. Forewing white shaded with ochreous, rectangular 
fuscous area along inner margin of middle third; two fuscous costal 
spots, one near middle and one at apical fourth; from latter a row of 
black dots extends from apex along termen to tornus; apical fourth 
flecked with black scales; two fuscous dots on cell, one basal, one 
distal; cilia brown basally and apically, white medially. Hindwing 
light fuscous with whitish costal margin; cilia light fuscous basally, 
white beyond. 

Male genitalia: Uncus arched, spatulate at apex; gnathos rounded 
at tip; vinculum complete, produced into dorsally projecting process 
in front; anellus with two laterally curved, pointed lobes; aedeagus 
long, slender; cornuti a cluster of heavy spines. 

Female genitalia: Genital plate small, undifferentiated. Ostium 
bursae sclerotized, opening at posterior margin of genital plate, 
pouchhke evagination between opening and inception of ductus 
seminalis; ductus bursae long, membranous; corpus bursae with 
large sclerotized signum; anterior apophyses short, not fused to 
genital plate. 

Type: South Fork of Cave Creek, Chiricahua Mts., Arizona. 
USNM 65824. 

Food plant: Unknown. 



42 PROCEEDINGS OF THE NATIONAL MUSEUM vol. hg 

Distribution: Arizona: South Fork of Cave Creek, Chiricahua 
Mts. (July) ; Madera Canyon, 4880 ft., Santa Kita Mts. (July). 

Described from the male holotype, July 4, 1939, collected by A. F. 
Braun, South Fork Cave Creek, Chiricahua Mts., Arizona; one male 
paratype, July 17, 1959, and four female paratypes, July 18, 19, 20, 
25, 1959, collected by Konald W. Hodges, Madera Canyon, 4880 ft., 
Santa Rita Mts., Arizona. 

This species is nearest A. haesitans, but it is readily separable by 
the vinculum produced into a dorsally projecting process in front 
in the male genitaUa and by the pouchlike evagination of the ostium 
bursae in the female genitalia. 

14. Antaeolricha vestalis (Zeller) 

Figures 9, 9a, 36; Plate 3b; Map 7 
Cryptolechia vestalis Zeller, 1873, Verh. Zool.-Bot. Ges. Wien, vol. 23, p. 247. 
Ide vestalis Walsingham, 1889, Insect Life, vol. 2, p. 155. 
Stenoma vestalis Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North America, 

vol. 4, p. 238. — Meyrick, 1922, in Genera insectorum, vol. 180, p. 82. 
Antaeolricha vestalis Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 30. 

Alar expanse 15-21 mm. 

Color: Face white; palpus white with dusky scales on exterior 
sides in male. Legs white sprinkled with dusky scales. Thorax 
white dorsally. Abdomen white. Forewing white, costal edge 
slightly ochreous. Hindwing white; cilia white. 

Male genitalia: Uncus curved, dilated at tip; gnathos large, divided 
into two distinct, sharp-pointed lobes; harpes as in A. schlaegeri; 
anellus with four lobes, ventral ones small and thumblike, bearing 
several setae at their apex, dorsal ones larger, blunter, bearing num- 
erous heavy setae. Aedeagus long, rounded at apex; cornuti absent. 

Female genitalia: Genital plate without distinct division into 
lamella ante vaginalis and lamella post vaginalis, ostium bursae small, 
opening near center of genital plate; ductus bursae long, membranous; 
corpus bursae with small, dentate signum ; inception of ductus seminalis 
near ostium. 

Type: In the Museum of Comparative Zoology, Harvard University. 

Type locality : Texas (C. vestalis) . 

Food plant: Unknown. 

Distribution: Florida: Lake Placid (April, Dec); Florida City 
(March) ; Royal Palm State Park (March, Sept.) ; Everglades (April) ; 
Fort Myers (April); Marco (April); St. Petersburg (Oct.); Paradise 
Key (March); Lakeland (March); Miami; Panacea (Aug., Oct.); 
Altamont; Glen wood. Georgia: Billy's Island, Okefenokee Swamp 
(June, July); Spring Creek (July). Mississippi: Ocean Springs 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 43 

(Aug.). SOUTH CAROLINA: DO locality (Feb.), texas: Burnet Co. 
(Oct.); Kerrville. new jersey: Lakehurst (July, Sept.). 

This species is similar to A. Irene but it is distinguishable by the 
apically dilated uncus, by the gnathos with two sharp-pointed lobes, 
and by the anellus with fom* lobes in the male genitalia. The absence 
of specialized structures around the ostium serves to distmguish the 
females. 

Walsingham (1889) syiionymized Harpalyce albella Chambers with 
I. vestalis; however, examination of Chambers' type located in the 
Museum of Comparative Zoology reveals that H. albella belongs in the 
genus Durrantia Busck and is so transferred. 

15. Antaeotricha manzanitae Keifer 

Figures 15, 37; Plate 3c; Map 7 
Antaeotricha manzanitae Keifer, 1937, California Dept. Agric. Bull., vol. 26, p. 334. 

Alar expanse 25-30 mm. 

Color: Face whitish ochreous; palpus overlaid with fuscous. Legs 
white, overlaid with fuscous; foreleg darkest, mid- and hindleg pro- 
gressively hghter. Thorax white dorsally with brownish-black tuft 
posteriorly. Abdomen white. Forewing white, irregularly shaded 
fuscous scales; two patches of dark scales at anal angle: apical margin 
with row of faint, transverse, fuscous dots. Cilia fuscous, white 
tipped. Hindwing light fuscous: cilia hghter. 

Male genitalia: Uncus curved, dilated at tip, terminating in two 
short spines: gnathos well developed, ending in short blunt tip; harpes 
as in A. schlaegeri; anellus with two lobes, one on each side of aedeagus. 
Aedeagus short and broad; cornuti a small cluster of spines. 

Female genitalia: Genital plate small, bandlike; ostium bursae and 
ductus bursae membranous: corpus bm'sae corrugated; inception of 
ductus seminalis near ostium. 

Type: In the California Academy of Science. 

Type locality: Shingle Springs, El Dorado Co., California. 

Food plant: Ardostaphylos sp. Keifer (1937) gives a thorough 
account of the life history of this species with numerous illustrations. 

Distribution : California : Shingle Springs, El Dorado Co. (April) ; 
Palm Desert (Apr.) ; Big Basin (July) ; Mt. Shasta City, Siskiyou Co. 
(July); Hat Creek, Shasta Co. (June). Oregon: Tiller (June). 
BRITISH Columbia: Wellington (July). 

This species is similar in size and color to A. schlaegeri, but it is 
separable by the presence of two small spines on the tip of the uncus 
in the male and by the cornjgated corpus bursae in the female. 



44 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Genus Stenoma Zeller 

Stenoma Zeller, 1839, Isis von Oken, vol. 32, p, 195. 

1. Stenoma mistrella Busck 

Figures 16, 39; Plate 3d; Map 8 

Stenoma mistrella Busck, 1907, Proc. Ent. See. Washington, vol. 8, p. 93. — 
Walsingham, 1913, Lepidoptera-Heterocera, vol. 4 (vol. 42 in Godman and 
Salvin, Biologla Central!- Americana), p. 183. — Barnes and Busck, 1920, 
Contrib. Nat. Hist. Lep. North America, vol. 4, p. 238. — Busck and Dampf, 
1929, Estud. Ofic. Fed. Agric. Mexico, vol. 2, p. 13.— Busck, 1934, in 
Lepidopterorum catalogus, vol. 67, p. 49. 

Alar expanse 20-23 mm. 

Color: Face white; palpus light fuscous. Legs ochreous; forelegs 
dark anteriorly. Thorax Ught ochreous dorsally. Abdomen dark 
fuscous. Forewing ranging from Hght ochreous, minutely speckled 
and overlaid with darker brown scales, to uniform dark fuscous; 
small, conspicuous, round black dot at apex of cell; costa nearly 
straight, apex pointed. Hindwing dark fuscous; ciha light ochreous. 

Male genitaha: Uncus reduced, knoblike; gnathos incomplete in 
front; harpes simple, somewhat pointed at apex, bearing short, stiff, 
bifurcate setae; vinculum complete; anellar lobes broad, pointed 
apicaUy; aedeagus large, flared apically; cornuti two clusters of very 
heavy spines. 

Female genitaha: Genital plate small, lamella postvaginahs fused 
medially with eighth sternite; ostium bursae large, somewhat sclero- 
tized; ductus bursae membranous; corpus bursae with Hghtly scle- 
rotized signum. Inception of ductus seminalis near ostium. 

Type: In the United States National Museum. 

Type locaUty: St. Louis, Missouri. 

Food plant: Phleum pratense L., bromegrass, and Kentucky blue- 
grass. Miller (1940) reports that the larvae construct sheltered tubes 
at the base of gxass similar to Crambus sp. and feed on the edge of 
the neighboring grass blades. 

Distribution: Mississippi: Jackson (July, Sept.); Pearl (June, 
Sept.). MISSOURI: St. Louis (April, July), ohio: Cincinnati (July). 
ILLINOIS: Chicago (June); Decatur (June, Aug., Sept.); Lacon (Sept.). 
TEXAS: Victoria (July), kansas: Onaga. Pennsylvania: Pittsburgh 
(Sept.). NEW MEXICO: Frijoles Canyon (Sept.). Manitoba: Win- 
nipeg. 

This species is readily separated from Stenoma crambitella by the 
flared apex of the aedeagus in the male and by the presence of only 
one lightly sclerotized signum in the female. 



NORTH AIVIERICAN STENOMIDAE — DUCKWORTH 45 

2. Stenoma crambitella Walsingham 

Figures 17, 40; Plate 3e; Map 8 

Stenoma crambitella Walsingham, 1889, Insect Life, vol. 2, p. 154. — Barnes and 
Busck, 1920, Contrib. Nat. Hist. Lep. North America, vol. 4, p. 238. — 
Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 38. 

Alar expanse 22 mm. 

Color: Face and palpus white, palpus shaded with pale brownish 
ochreous scales. Legs whitish, shaded with brown scales. Thorax 
white dorsally. Abdomen whitish ochreous. Forewing white, few 
grayish fuscous scales on extreme costal margin basally and single 
dot of same color at apex of cell, apex somewhat pointed. Hindwing 
white, tinged with ochreous; cilia white. 

Male genitalia: Uncus reduced in size, but not as much as in S. 
mistrella; gnathos complete in front, forming a band; harpes as in S. 
mistrella, but blunt at apex; vinculum complete; anellar lobes long, 
slender; aedeagus short, broad; cornuti a small cluster of heavy spines. 

Female genitalia: Genital plate small, lamella postvaginalis fused 
laterally with eighth sternite ; ostium bursae large ; ductus bursae long, 
membranous; corpus bursae with two lightly sclerotized, dentate 
signa. Inception of ductus seminalis near ostium. 

Type: In the British Museum (Natural History). 

Type locality: Arizona. 

Food plant: Unknown. 

Distribution: Arizona: Palmerlee; Redington; Santa Rita Mts. 
(June); Santa Catalina Mts.; Chiricahua Mts. (Aug.). texas: 
Kerrville (June). 

This species differs from Stenoma mistrella in the differently shaped 
aedeagus in the male and in the presence of two lightly sclerotized 
signa in the female. 

Genus Setiostoma Zeller 

Setiostoma Zeller, 1875, Verh. Zool.-Bot. Ges. Wien, vol. 25, p. 324. 

1. Setiostoma xanthobasis Zeller 

Figures 18, 18a, 41; Plate 3f; Map 9 

Setiostoma xanthobasis Zeller, 1875, Verh. Zool.-Bot. Ges. Wien, vol. 25, p. 324. — 
Busck, 1925, Proc. Ent. Soc. Wasliington, vol. 27, p. 48; 1934, in Lepidop- 
terorum catalogus, vol. 67, p. 2. 

Alar expanse 12-14 mm. 

Color: Face and palpus lemon yellow. Legs dark brown; foreleg 
with coxa entirely white, tarsi ringed with white; midleg with two 
tibial rings; tibial spurs and tarsal rings white. Thorax brown 



46 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

dorsally except tegulae, which are lemon yellow. Abdomen brown 
with patches of white scales laterally. Forewing deep brown, with 
lemon yellow triangular area, continuous with yellow tegula, occupying 
basal third except short costal lenticular spot and longer, narrower 
spot along posterior wing margin, both spots concolorous with ground 
color. Apical two-thirds of forewing with inconspicuous area of white 
scales near midlength of costal margin and similar, smaller area 
slightly more distad; broad transverse band, parallel to distal margin 
of yellow spot, small group of scales behind more basal white area, 
short curved band extending caudolaterad from more distal white 
area, submarginal longer straight band parallel to outer wing margin, 
iridescent blue. Cilia blackish brown. Hindwing dark brown with 
patch of white on basal half of anterior margin ; cilia brown edged in 
white. 

Male genitalia: Uncus pointed; gnathos a simple band; harpes 
simple with palmate multilobed hairs on outer part of costa; vinculum 
narrow, incomplete in front, anellus with two flattened, upright lobes; 
aedeagus large with apex pointed; cornuti a large cluster of small 
spines and one large single spine. 

Female genitalia: Genital plate membranous; ostium bursae large, 
trumpet-shaped; ductus bursae membranous; corpus bursae with 
signum large, dentate; inception of ductus seminalis near ostium. 

Type : In the Museum of Comparative Zoology, Harvard University. 

Type locality: Texas. 

Food plant: Quercus sp. According to Forbes (1923), the larva 
is found in a nest about two centimeters in diameter, formed by an 
oval wall of silk between two sUghtly separated oak leaves, and it 
feeds on the lower parenchyma only. 

Distribution: Massachusetts: Martha's Vineyard, new jersey: 
Lacy (July). Maryland: Annapohs (Aug.). district of Columbia: 
Washington (July), north Carolina: Southern Pines (Aug.). 
Georgia: Spring Creek (July). Florida: Enterprise (April); Lake- 
land (May). Illinois: no locality. Missouri: no locality (May). 

This species is very closely related to S. Jernalddla Riley, but it 
is distinguished readily by its single large spine and by a cluster of 
small spines in the aedeagus of the male, and by its heavily sclero- 
tized, dentate signum in the corpus bursae of the female. 

2. Setiostonia fernaldella Riley 

Figures 19, 19a, 42; Plate 4a; Map 9 

Setiostonia fernaldella Riley, 1889, Proc. Ent. Soc. Washington, vol. 1, p. 155. — 
Busck, 1934, in Lepidopterum catalogus, vol. 67, p. 1. 

Alar expanse 12-13 mm. 

Color: Face yellow sprinkled with brown; palpus yellow. Legs 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 47 

blackish brown ; foreleg with coxa white or yellowish, tarsi ringed with 
white; spurs of midtibia white, with few dusky scales, and mid- 
tarsi ringed with white ; hindleg with two tibial rings, spurs and tarsal 
rings white. Thorax greenish yellow dorsally sprinkled with brown 
scales. Abdomen dark brown with metallic iridescence. Forewing 
with triangular marking of basal third as in S. xanthobasis but greenish 
yellow, concolorous and continuous with greenish yellow of thorax, 
and with darker spot along posterior margin much shorter and more 
restricted to anal region. Apical two-thirds dark brown with iri- 
descent bronze-to-violet markings, in reflected light, in transverse 
band across middle third of wing and an area that curves near wing 
apex and extends parallel to outer wing margin. Hindwing as in 
S. xanthobasis. 

Male genitalia: As in S. xanthobasis with exception of anellus, 
which bears upright, median, dentate process in addition to two 
lateral flattened upright processes, and cornuti, which consist of 
several large single spines. 

Female genitalia: As in S. xanthobasis with exception of absence 
of signum in corpus bursae and presence of sclerotized plate at junction 
of ostium bursae and ductus bursae. 

Type: In the United States National Museum. 

Type locality: Los Angeles, California. 

Food plant: Quercus wislizenii A. DeCandoUe and Quercus agri- 
folia Nee. 

Distribution: California: Forest Home, San Bernardino Co. 
(June) ; Los Angeles Co. (July) ; San Diego (June, July) ; Baldy Mts. 
(June); Pasadena (June); Santa Clara. Arizona: Madera Canyon, 
Santa Rita Mts. (July, Aug., Sept.); Pena Blanca Canyon (Sept.). 

This species is readily distinguished from S. xanthobasis by its 
cornuti that consists of more than one large single spine in the 
aedeagus of the male and by the absence of a signum in the corpus 
bursae of the female. 

Genus Mothonica Walsingham 

Mothonica Walsingham, 1912, Lepidoptera-Heterocera, vol. 4 (vol. 42 in Godman 
and Salvin, Biologia Centrali- Americana), p. 153. 

1. Mothonica kimballi, new species 

Figures 20, 20a, 38; Plate 4b; Map 10 

Alar expanse 17-19 mm. 

Face ochreous shaded with fuscous; palpus white sprinkled with 
fuscous, apical segment tipped with fuscous, second segment shaded 
with fuscous basally. Legs creamy white, hindtibia with long hairs 
above, fore- and midleg shaded heavily with fuscous. Thorax 

728-597—84 3 



48 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

smooth, white sprmkled with fuscous dorsally. Abdomen white, 
shaded to various degrees with fuscous. Forewing long, somewhat 
narrow; ground color white, with three large fuscous areas along costa; 
marginal and submarginal transverse row of fuscous dots at apex, 
marginal row darker and giving sinuated effect to termen; another 
area of dark fuscous raised scales at anal angle. Anterior 
half of cilia light fuscous, posterior half white. Hindwing very Hght 
fuscous; cilia fuscous basally, white apically. 

Male genitalia: Uncus simple, short, with group of setae at apex; 
gnathos two lateral plates flanking tuba analis; harpes with pincer- 
shaped costa, apex somewhat pointed and recurved, bearing heavy 
bifurcate setae; sacculus with prominent bifurcate lobe; vinculum 
complete; anellus a large rectangular plate with large upright lateral 
lobes; aedeagus large basally, tapering apically to a point, cornuti two 
large clusters of heavy spines. 

Female genitalia: Genital plate large, with large median slot. 
Ostium bursae sclerotized, ductus bursae short, membranous; corpus 
bursae large, with large dumbbell-shaped, dentate signum; anterior 
apophyses long, not fused to genital plate. 

Type: Siesta Key, Sarasota County, Florida. USNM 65825. 

Food plant: Unknown. 

Distribution: Florida: Siesta Key, Sarasota County (Jan., Feb., 
March, April, May, Nov.); Vero Beach (April). 

Described from the male holotype, Jan. 24, 1954, two male para- 
types, Feb. 26, 1954, Jan. 15, 1954, collected by C. P. Kimball, 
Siesta Key, Sarasota County, Florida; two male paratypes, April 2, 
1941, collected by J. R. Malloch, Vero Beach, Florida; one female 
paratype, March 4, 1953, collected by C. P. Kimball, Siesta Key, 
Sarasota County, Florida. 

This is the first species of the genus Mothonica to be described from 
North America. It is very similar to Mothonica fluminata (Meyrick), 
a Colombian species. The bifurcate lobe on the harpe of M. kimballi 
readily separates the males. No females of M. fluminata were 
available for comparison. 

This species is named in honor of Mr. C. P. Kimball of West 
Barnstable, Massachusetts, who provided material from his personal 
collection for description. 

Genus Menestomorpha Walsingham 

Menestomorpha Walsingham, 1907, Proc. U.S. Nat. Mus., vol. 33, p. 214. 

1. Menestomorpha oblongata Walsingham 

Figures 21, 43; Plate 4c; Map 11 
Menestomorpha oblongata Walsingham, 1907, Proc. U.S. Nat. Mus., vol. 33, 
p. 215. — Barnes and Busck, 1920, Contrib. Nat. Hist. Lep. North America, 
vol. 4, p. 238. — Busck, 1934, in Lepidopterorum catalogus, vol. 67, p. 4. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 49 

Alar expanse 15 mm. 

Color: Face and palpus white flecked with brown scales, palpus 
brown at tip. Legs white flecked with brown scales, tarsi ringed 
with brown. Thorax white flecked with brown scales. Abdomen 
with mixed brown and white scales. Forewing white with two trans- 
verse brown bands converging at anal angle forming V-shaped mark 
on basal third, middle third with indistinct brown shading blending 
into indistinct brown streaks in apical third that follow lines of veins 
beyond cell to termen and costa; with row of five or six indistinct 
brownish dots along termen reaching to apex; cilia brown tipped in 
white. Hindwing light brown; cilia lighter, with light greyish-fuscous 
line along margin at base and two parallel shades running through it. 

Male genitalia: Uncus very small bearing several small setae at 
apex; gnathos with two small processes beneath uncus; harpes with 
large lobe basally, median forked structure, and somewhat pointed 
apical lobe bearing short, bifurcate setae; vinculum complete; anellus 
with two large, pointed processes surrounding and extending above 
aedeagus; aedeagus short, pointed laterally; cornuti a small cluster of 
heavy spines. 

Female genitalia : Genital plate reduced to simple band surrounding 
ostium; ostium bursae and ductus bursae membranous; corpus 
bursae large; membranous, with two dentate, lightly sclerotized signa. 

Type: In the British Museum (Natural History). 

Type locality: Fort Grant, Arizona. 

Food plant: The type series was reared from a cynipid gall on 
Quercus sp. 

Distribution: Arizona: Fort Grant (April); Baboquivari Mts. 
(Oct.); Madera Canyon, Santa Rita Mts. (July, Aug., Sept., Oct.). 
caldfornia: San Bernardino Co. (June). 

Genus Menesta Clemens 

Menesta Clemens, 1860, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, p. 213. 
1. Menesta tortriciformella Clemens 

Figures 23, 44; Plate 4d; Map 12 

Menesta tortriciformella Clemens, 1860, Proc. Acad. Nat. Sci. Philadelphia, 
vol. 12, p. 213; 1872, The Tineina of North America, p. 151. — Chambers, 
1878, Bull. U.S. Geol. Surv., vol. 4, p. 157.— Walsingham, 1881, Proc. Zool. 
Soc. Loudon, p. 319; 1889, Insect Life, vol. 2, p. 154.— Busck, 1903b, Proc. 
U.S. Nat. Mus., vol. 25, p. 903; 1934, in Lepidopterorum catalogus, vol. 
67, p. 5. 

Gelechia liturella Walker, 1864, List of the specimens of lepidopterous insects in 
the collection of the British Museum, vol. 29, p. 591. 

Hyale coryliella Chambers, 1875, Cincinnati Quart. Journ. Sci., vol. 2, p. 242. 

Strobisia albaciliaeella Chambers, 1878, Canadian Ent., vol. 10, p. 77. (New- 
synonymy.) 



50 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Menesta albaciliaeella Busck, 1903b, Proc. U.S. Nat. Mus., vol. 25, p. 903; 1934, 

in Lepidopterorum catalogus, vol. 67, p. 5. 
Menesta albiciliella [sic] Walsingham, 1911, Lepidoptera-Heterocera, vol. 4 

(vol. 42 in Godman and Salvin, Biologia Central!- Americana) , p. 104. 
Menesta albaciliella [sic] Braun, 1915, Ent. News, vol. 26, p. 160. 

Alar expanse 9-10 mm. 

Color: Face and palpus white, palpus shaded with fuscous. Legs 
white, shaded anteriorly with fuscous, especially apex of tibia and 
apices of tarsi, hindtibia with long haii-s. Thorax brown doi-sally, 
with gi-ayish hue. Abdomen brown dorsally, white ventrally. Fore- 
wing blackish brown with gi'eenish-violet reflections; few white scales 
just beyond middle forming indistinct, short, transverse line; apical 
cUia white or brown, remainder brown. Hindwing much lighter 
brown; cUia brown, tipped with white apically. 

Male genitalia: Uncus simple, large basally, tapering to point 
apically, curved ventrad parallel to gnathos; gnathos well developed, 
very long, blunt apically; harpes simple, bluntly rounded at apex, 
bearing large cluster of small, stiff, bifurcate setae; vinculum complete, 
anellus with two lateral lobes partially encircling aedeagus, aedeagus 
pointed apically; cornuti consisting of two large, heavy spines. 

Female genitalia: Genital plate reduced to small median triangular 
sclerite; ostium bursae slightly sclerotized; ductus bursae with dilated 
area before inception of ductus seminalis; corpus bm-sae with large, 
heavily sclerotized, cross-shaped signum with outwardly projecting 
median lobe; inception of ductus seminalis approximately halfway 
between ostium and corpus bursae. Anterior apophyses free, arising 
ventrally. 

Type: In the Academy of Natural Sciences of Philadelphia. 

Type locality: Pennsylvania (M. tortriciformella), Kentucky (H. 
coryliella), Nova Scotia (0. litur ella) ,Cmcmnsiti, Ohio (S. albaciliaeella). 

Food plant: Rubus Villosus Ait. and Corylus americana Walt. 
Chambers (1875) and Braun (1915) give accounts of the life history of 
this species. 

Distribution: ohio: Cincinnati (May, June, July). Pennsylvania: 
Harvey's Lake (June), new jersey: Essex Co. (June), new york: 
Buffalo (June); Peru (June); Protection (June). Maryland: Cabin 
John (May). Virginia: Ocean View (July). Ontario: Kearney 
(July). 

This species has been known previously only from its type, which 
Busck (1903) incorrectly reported as being without an abdomen. 
The genitalia of the type of M. tortriciformella has been studied and 
found to be identical with those of S. albaciliaeella. This synonymy 
is not surprising because the only characters used in the past to 
separate the two species were slight differences in the color of the head 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 51 

and the presence or absence of white apical cUia on the forewing. 
This species closely resembles M. melanella in both superficial 
appearance and in structure of the genitalia; however, M. tortrici- 
formella lacks on the costa of the forewing the white spot that is 
present in M. melanella. The genitalia differ from those of AI. melan- 
ella in that the cornuti in the aedeagus of the male consist of two large 
heavy spines and the signum in the corpus bursae of the female is 
cruciform. 

2. Menesta melanella Murtfeldt 

Figures 22, 45; Plate 4e; Map 12 
Menesta melanella Murtfeldt, 1890, Insect Life, vol. 2, p. 304. — Busck, 1903b, 
Proc. U.S. Nat. Mus., vol. 25, p. 903; 1934, in Lepidopterorum catalogus, 
vol. 67, p. 5. 

Alar expanse 10-12 mm. 

Color: Face and palpus white. Legs white shaded lightly with 
fuscous. Thorax blackish brown dorsally. Abdomen blackish brown 
dorsally, white ventrally. Forewing blackish brown with greenish- 
violet reflections, triangular patch of white scales on costa about mid- 
way between base and apex; ciha brownish black, white apically. 
Hindwing with broad white streak extending along costa from base 
to beyond midpoint; cilia brown with patches of white near outer 
angle and near base. 

Male genitalia: Uncus as in AI. tortriciformella except apex not as 
sharply pointed; gnathos as in M. tortriciformella except apex, which 
ends in a sharp point; harpes and vinculum as in M. tortriciformella; 
anellus with two lateral lobes as in M. tortriciformella but larger and not 
encircling aedeagus; aedeagus slightly pointed apically; cornuti con- 
sisting of one large heavy spine. 

Female genitaha: Genital plate, ostium bursae, and ductus bursae 
as in M. tortriciformella; corpus bursae with heavily sclerotized, rec- 
tangular signum with median lobe; inception of ductus seminalis 
near ostium, anterior apophyses as in M. tortriciformella. 

Type: In the United States National Museum. 

Type locality: Missouri. 

Food plant: Quercus stellata Wangh. Murtfeldt (1890) gives an 
account of the life history of this species. 

Distribution: south Carolina: Oconee (Aug.). new jersey: Lake- 
hurst (June). Massachusetts: Martha's Vineyard (June). Florida: 
Dade City (April). Virginia: Falls Church (Aug.). Arizona: Ma- 
dera Canyon, Santa Rita Mts. (Aug.). 

From M. tortriciformella, which it resembles, this species differs by 
a white spot being present on the costal edge of the forewing, by the 
cornuti consisting of one large heavy spine in the male genitalia, and 
by the signum being rectangular in the female genitaha. 



52 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Literature Cited 

Barnes, W., and Busck, A. 

1920. Notes and new species. Contrib. Nat. Hist. Lep. North America, 
vol. 4, pp. 236-240, 256-258. 
Braun, a. F. 

1915. Life history of Menesta albaciliella Chambers (Lep.). Ent. News, 
vol. 26, pp. 160-161. 
BuscK, A. 

1903a. Notes on Brackenridge Clemens' types of Tineina. Proc. Ent. Soc. 

Washington, vol. 5, pp. 181-232. 
1903b. A revision of the American moths of the family Gelechiidae, with 
descriptions of new species. Proc. U.S. Nat. Mus., vol. 25, pp. 
767-938. 

1907. New American Tineina. Proc. Ent. Soc. Washington, vol. 8, pp. 

86-99. 

1908. Two new stenomid moths from the Eastern United States. Proc. 

Ent. Soc. Washington, vol. 10, pp. 111-112. 
1921a. On the male genitalia of the Microlepidoptera and their systematic 

importance. Proc. Ent. Soc. Washington, vol. 23, pp. 145-152. 
1921b. Microlepidoptera from British Columbia. Canadian Ent., vol. 53, 

pp. 276-280. 
1925. On the genus Setiostoma Zeller (Lepidoptera: Stenomidae). Proc. 

Ent. Soc. Washington, vol. 27, pp. 48-50. 
1934. Stenomidae. In Lepidopterorum catalogus, vol. 67, pp. 1-73. 
BuscK, A., and Dampf, A. 

1929. Una palomilla {Stenoma crambtna Busck) como una nueva plaga del 

algodon en el Estado de Oaxaca. Estud. Ofic. Fed. Agric. Mexico, 

vol. 2, pp. 1-55. 
Chambers, V. T. 

1874. North American Microlepidoptera. Canadian Ent., vol. 6, pp. 

229-249. 

1875. Tineina of the United States. Cincinnati Quart. Journ. Sci., vol. 

2, pp. 226-259. 

1877. Tineina of Colorado. BuU. U.S. Geol. Surv., vol. 3, pp. 121-142. 

1878. Micro-Lepidoptera. Canadian Ent., vol. 10, pp. 74-78. 
Clarke, J. F. Gates 

1941. The preparation of slides of the genitalia of Lepidoptera. Bull. 

Brooklyn Ent. Soc, vol. 36, pp. 149-161. 
1955a. Catalogue of the type specimens of Microlepidoptera in the British 

Museum (Natural History) described by Edward Meyrick, vol. 1, 

332 pp., pis. 
1955b. Catalogue of the type specimens of Microlepidoptera in the British 

Museum (Natural History) described by Edward Meyrick, vol. 2, 

531 pp., pis. 
Clemens, B. 

1860. Contributions to American lepidopterology. Proc. Acad. Nat. Sci. 

Philadelphia, vol. 12, pp. 4-15, 156-174, 203-221, 345-362, 

522-547. 
1863. American Micro-Lepidoptera. Proc. Ent. Soc. Philadelphia, vol. 2, 

pp. 4-14, 119-129. 
1872. The Tineina of North America, compil. H. T. Stainton. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 53 

Fletcher, T. B. 

1929. A list of the generic names used for Micro lepidoptera. Mem. Dept. 
Agric. India, Ent. Ser., vol. 11, pp. 1-244. 
Forbes, W. T. M. 

1923. The Lepidoptera of New York and neighboring states. Cornell 
Agric. Exp. Sta. Mem., vol. 68, pp. 3-729. 
Frost, S. W. 

1931. The well-marked fruit-worm, Stenoma algidella Walk. Journ. Econ. 

Ent., vol. 24, pp. 1208-1213. 
Heinrich, C. 

1921. Some Lepidoptera likely to be confused with the pink boUworm. 

Journ. Agric. Res., vol. 20, pp. 807-836. 
Janse, a. J. T. 

1932. The moths of South Africa, vol. 1, 376 pp., 15 pis. 
Keifer, H. H. 

1937. California Microlepidoptera, XII. California Dept. Agric. Bull., vol. 

26, pp. 334-338. 
Meyrick, E. 

1880. Descriptions of Australian Microlepidoptera, IV: Tineina (continued). 

Proc. Linn. Soc. New South Wales, vol. 5, pp. 204-271. 
1883. On the classification of some families of the Tineina. Trans. Ent. 

Soc. London, pp. 119-131. 

1889. On the interpretation of neural structure. Ent. Monthl. Mag., ser. 1, 

vol. 25, pp. 175-178. 

1890. Descriptions of Australian Lepidoptera, Xylorctidae. Trans. Roy. 

Soc. South Australia, vol. 13, p. 23. 

1906. Descriptions of Australian Tineina. Trans. Roy. Soc. South Aus- 
tralia, vol. 30, pp. 33-66. 

1909. Descriptions of Microlepidoptera from Bolivia and Peru. Trans. 
Ent. Soc. London, pp. 13-43. 

1912. Descriptions of South American Microlepidoptera. Trans. Ent. Soc. 

London, pp. 673-718. 

1913. Descriptions of South American Microlepidoptera. Trans. Ent. Lon- 

don, pp. 170-200. 
1915-1934. Exotic Microlepidoptera, 4 vols. 

1922. Oecophoridae. In Genera insectorum, vol. 180, pp. 1-224. 

1928. A revised handbook of the British Lepidoptera. 

1929. The Microlepidoptera of the "St. George" expedition. Trans. Ent, 

Soc. London, vol. 76, pp. 489-521. 

1930. Ergebnisse einer Zoologischen Sammelreise nach Brasilien, insbeson- 

dere in das Amazonasgebiet, ausgefuhrt von Dr. H. Zerny V. Teil: 
Microlepidoptera. Ann. Naturhist. Mus. Wien, vol. 44, pp. 223- 
268. 

1931. Microlepidoptera from South Chile and Argentina. Anal. Mus. Nac. 

Hist. Nat. Buenos Aires, vol. 36, pp. 377-415. 
Miller, H. D. O. 

1940. Observations on Stenoma mistrella Busck. Journ. Kansas Ent. Soc, 
vol. 13, pp. 1-3. 
Murtfeldt, M. E. 

1890. An interesting Tineid. Insect Life, vol. 2, pp. 303-305. 
RiLET, C. V. 

1889. Two brilliant and interesting Micro-Lepidoptera new to our fauna. 
Proc. Ent. Soc. Washington, vol. 1, pp. 155-159. 



54 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

TUXEN, S. L. 

1956. Taxonomist's glossary of genitalia in insects. 
Walker, R. 

18G4. List of the specimens of lepidopterous insects in the collection of the 
British Museum, pt. 29. 
Walsingham, L. 

1881. On some North American Tineidae. Proc. Zool. Soc. London, pp. 

301-325. 
1889. Steps towards a revision of Chambers' index, with notes and descrip- 
tions of new species. Insect Life, vol. 2, pp. 150-155. 
1907. Descriptions of new North American Tineid moths, with a generic 
table of the family Blastobasidae. Proc. U.S. Nat. Mus., vol. 33, 
pp. 197-228. 
1911.-1913. Lepidoptera-Heterocera, vol. 4 (vol. 42 in Godman and 
Salvin, Biologia Centrali-Americana), pp. 49-112 (1911); 113-118 
(1912); 169-190 (1913). 
Zeller, p. C. 

1839. Versuch einer naturgemassen Eintheilungun der schaven: Tinea. 
Isis von Oken, vol. 32, pp. 167-222. 

1854. Die Depressarien und einige ihnen nahe stechenden Gattungen. Linn. 

Entom., vol. 9, pp. 189-403. 

1855. Nachtrag zu den im 9. Bande der Linnaea Ent. beschriebenen arten 

des genus Cryptolechia. Linn. Entom., vol. 10, pp. 145-168. 
1873. Beitrage zur Kenntniss der nordamericanischen Nachtfaltern, besond- 

ers der Microlepidopteren, part 2. Verh. Zool.-Bot. Ges. Wien, 

vol. 23, pp. 201-334. 
1875. Beitrage zur Kenntniss der nordamericanischen Nachtfaltern, besond- 

ers der Microlepidopteren, part 3. Verh. Zool.-Bot. Ges. Wien, 

vol. 25, pp. 207-360. 



PROC. U.S. NAT. MUS. VOL 116 



DUCKWORTH PLATE 1 




Pl,\ti-; 1. — a, Antaeolricha ichlaegeri Zcller; b, A. lindseyi (Barnes and Busck); c, ./. 
unipunctt'lla (Clemens); d, A. leucillona (Zcller); e, A. nsseella (Walsinghani); 
f. A. decorosella (Busck). 



PROC. US NAT. MUS. VOL 116 



DUCKWORTH— PLATE 2 




Plate 2. — a, Antaeutricha furcata (VValsiiiyhaiii); b, A. irfiw (Barnes and P>usck); c, 
A. humilis (ZcUer); d, A. agrioschista (Mcyrick); c, A. tli'niiasi (Barnes and Busck); 
f, A. haesitans (Walsingham). 



PROC US NAT. MUS. VOL 116 



OUCKWORTH PLATE 3 




Plate ,v — a, Antaeotricha juscorectangulata, new species; b, A. vestalis (Zeller); 
c, A. manaznitae Keifer; d, Stenoma mistrella Busck; e, S. crambitella VValsingham; 
f, Setiostoma xanthobasis Zellcr. 



PROC. U.S. NAT. MUS. VOL. 116 



DUCKWORTH — PLATE 4 




Plate 4. — a, Seliostotna fernuldella Rile}"; b, Mothunica kiniballi, new species; c, 
Menestomorpha oblongata W'alsiiiKham; d. Mt-nesta torlricifonnella Clemens; 
e, M. melanella Murtfcldt. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 



55 







f^-^.rA i 



I •• Anlaeolrlcha schloegerKZellar) , 

I ▲ Anloeotrlcho lindstyi (Barnes a Busch) 




Map 1. — Distribution of Antaeotricha schloegeri (Zeller) and A. lindseyi (Barnes and Busck). 




^^^^ 




ft--^-^ i 



/• V 



Kr-\ 



• Anloeotrlcho wtipunct«Uo (Clement] 




Map 2. — Distribution of Antaeotricha unipuncUlla (Clemens). 



726-597—64 5 



56 



PROCEEDENTGS OF THE NATIONAL MUSEUM 





/ ;. 






h--- t--v 



^ 



vr' 



AnloeotrlchaJ«uclllano (Zeller) 



Map 3. — Distribution of Antaeotricha leucillana (Zeller). 



IPf^-C T 



/ I 



\^M 



• AntQsolricho osseello (Wctsinghom) 
A Antaeotricha decorosello (BuscK) 

* Anioeolricho furcota (WaUlngliam) 




Map 4. — Distribution of Antaeotricha osseella (Walsingham), A. decorosella (Busck), and 

A. fuTcata (Walsingham). 



NORTH AMERICAN STENOMIDAE DUCKWORTH 



57 



[^M-^ 



• Aniaeolrlcha iren* (Sarnes A Busck) 
A Antoeotncha humISs lZe(I£r) 

* Antoeovicha optenMita (Me/rick) 




Map S. — Distribution of Antaeotricha Irene (Barnes and Busck), A. humilis (Zeller), and 

A. agrioschista (Meyrick). 





/ \.-. 



• Aiila«otricho thoma^ (Barnts 6 Buick) 
▲ AnloeotrtclM ho«sitan< (Woluofhom) 



Map 6. — Distribution of Antaeotricha thomasi (Barnes and Busck) and A. haesitans (Wal- 

singham). 



58 



PROCEEDINGS OF THE NATIONAL MUSEUM 



vot. 116 




Map 7. — Distribution of Antaeotricha fuscorectangulata, new species, A. vestalis (Zeller), 

and A. manzanitae (Keifer). 




Map 8. — Distribution of Stenoma mistrella (Busck) and S. cramhitella (Walsingham). 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 



59 



m--^_ \ T 





]-■- M ( 1 \ /-V"^ 



I 



\/'-\ 



• Setiottomo lanthobasis (Zeller) 
▲ Stttostoma fernold«1la (Ritey) 




Map 9. — Distribution of Setiostoma xanthobasis (Zeller) and S. fernaldella (Riley). 




Map 10. — Distribution of Mothonica kimballi, new species. 



60 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 





• Menestomorpha oblongoto {WolsJnghom} 



Map 11. — Distribution of Menestomorpha oblongata (Walsingham). 





"--^■•J.j-'-'i 1 



• Menesia tortriciformelta (Clemens) 
A Menesta melonella (Murtfeldt) 



Map 12. — Distribution of Menesta tortriciformella (Clemens) and M. melanella (Murtfeldt). 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 



61 




Figures 1-5. — Ventral view of male genitalia: 1, Antaeotricha schlaegeri (Zeller); 2, A. 
lindseyi (Barnes and Busck); 3, A. unipunculla (Clemens); 4, A. leucillana (Zeller); 5, A. 
osseella (Walsingham). 



62 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 6-9. — Ventral view of male genitalia: 6, Antaeotricha decorosella (Busck); 7, A. 
fuTcata (Walsingham); 8, A. irene (Barnes and Busck); 9, A. vestalis (Zeller). Lateral 
view of aedeagus: 8a, A. irene; 9a, A. vestalis. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 



63 




Figures 10-14. — Ventral view of male genitalia: 10, Antaeotricha agrioschista (Meyrick); 
II, A. humilis (Zeller); 12, A. thomasi (Barnes and Busck); 13, A. haesitans (Walsingham); 
14, A. fuscorectangulata, new species. Lateral view of aedeagus: 10a, A. agrioschista; 
14a, A. fuscorectangulata. 
726-597—64 4 



64 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 15-17. — V«itral view of male genitalia: 15, Antaeotricha manzanitae Keifer; 
16, Stenoma mistrella Busck; 17, S. cramhitella Walsingham. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 



65 




Figures 18-21. — Ventral view of male genitalia: 18, Setiostoma xanthobasis Zeller; 19, S. 
fernalddla Riley; 20, Mothonica kimballi, new species; 21, Menestomorpha oblongata 
Walsingham. Lateral view of aedeagus: 18a, S. xanthobasis; 19a, S. fernaliella; 20a, M. 

kimballi. 



66 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. iis 




Figures 22-26.— Ventral view of male genitalia: 22, Menesta melanella Murtfeldt; 23, M. 
tortriciformella Clemens. Ventral view of female genitalia: 24, Antaeotricha schlaegeri 
(Zeller); 25, A. unipunctella (Clemens); 26, A. leucillana (Zeller), 



NORTH AMERICAN STEXOMIDAE — DUCKWORTH 



67 




Figures 27-30. — Ventral view of female genitalia: 27, Antaeotricha osseella (Walsingham); 
28, A.furcata (Walsingham); 29, A. irene (Barnes and Busck); 30, A. decorosella (Busck). 



68 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 31-34.— Ventral view of female genitalia: 31, Antaeotricha humilis (Zeller); 32, A. 
thomasi (Barnes and Busck); 33, A. fuscorectangulata, new species; 34, A. agrioschista 
(Meyrick). 



NORTH AMERICAN STENOMIDAE DUCKWORTH 



69 




Figures 35-37. — Ventral view of female genitalia: 35, Antaeotricha haesitans (Walsingham); 
36, A. vestalis (Zeller); 37, A, manzanitae Keifer. 



70 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 38-41. — Ventral view of female genitalia: 38, Moihonica kimballi, new species; 
39, Stenoma mistrella Busck; 40, S. crambitella Walsingham; 41, Setiostoma xanthobasis 
Zeller. 



NORTH AMERICAN STENOMIDAE — DUCKWORTH 



71 




Figures 42^5. — Ventral view of female genitalia: 42, Setiostoma fernaldella Riley; 43, 
Menestomorpha oblongata Walsingham; 44, Menesta tortriciformella Clemens; 45, M. 
melanella Murtfeldt. 



Proceedings of ^^^ 

the United States R^^l§ 
National Museum ^^^0^ 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1964 Number 3496 

A SURVEY OF VERTEBRAL NUMBERS IN SHARKS > 



By Victor G. Springer and J. A. F. Garrick 



Introduction 



This paper broadly surveys vertebral numbers in sharks. The 
study was prompted by our discovery that vertebral numbers are 
important systematic characters in those carcharinid shark genera 
that we have been investigating (Springer, 1964; Garrick, in ms). 
We, therefore, have undertaken to determine if vertebral numbers 
are of similar value in other genera, with the hope that some 
contribution might be made to shark classification as a whole. 

Vertebral numbers have not been used previously as a systematic 
character in sharte although they have received some attention in 
rays (Ishiyama, 1958) and have been widely employed in teleosts 
(e.g., BaUey and Gosline, 1955 ; Schmidt, 1917). The vertebral numbers 
from sharlvs th&t have been recorded in the literature are given either 
without comment or comparison or are employed as data for studies 
in morphology or intraspecific variation (Punnett, 1904; Aasen, 1961). 

We present here vertebral data on 1524 specimens. We personally 
made counts from 858 of these, mostly by X-ray methods. The 



• This study was conducted on funds administered by Dr. Leonard P. Schultz and provided by contracts 
between the Office of Naval Research (NONR 1354(09)), the Atomic Energy Commission (AEC AT(30-1)- 
2400), and the Smithsonian Institution. 

73 



74 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

remaining 666 counts are from the literature or were supplied by 
colleagues. The 1524 specimens pertain to 70 of the approximately 
80 genera of sharks and to 135 of the approximately 300 species. 
Because our purpose has been to survey', our coverage within in- 
dividual species is far from complete, but considerable attention has 
been paid to a few species that have presented problems. 

We wish to express our gratitude to the officials of the following 
institutions for their assistance in providing specimens and X-ray 
facilities. 

ANSP — Academy of Natural Sciences of Philadelphia 

BMNH — British Museum (Natural History), London 

CAS — Cahfornia Academy of Sciences, San Francisco 

CM — Canterbury Museum, Christchurch, New Zealand 

CNHM — Cliicago Natural History Museum 

DMNZ — Dominion Museum, Wellington, New Zealand 

GVF — George Vanderbilt Foundation, Stanford University, California 

IRSN — Institut Royal des Sciences Naturelles de Belgique, Brussels 

ISZZ — Institut fiir Spezielle Zoologie und Zoologisches Museum, BerUn 

MCSN — Museo Civico di Storia Naturale, Genoa, Italy 

MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, 

Massachusetts 
RNH — Rijksmuseum van Natuurlijke Historic, Leiden, Holland 
SIO — Scripps Institution of Oceanography, La Jolla, California 
SU — Division of Systematic Biology, Stanford University, California 
TGFC — Texas Game and Fish Commission Marine Laboratory-, Rockport 
UBC — University of British Columbia, Vancouver 

UMML — University of Miami Institute of Marine Sciences, Miami, Florida 
UMMZ — University of Michigan Museum of Zoology, Ann Arbor 
USNM — United States National Museum, Wasliing-ton, D.C. 
UZMK — Universitetets Zoologiske Museum, Copenhagen, Denmark 

J. G. Casey, Bureau of Sport Fisheries, Sandy Hook Marine Labora- 
tory, kindly provided some vertebral counts of Alopias superciliosus 
and Isurus oxyrinchus, and H. Heyamoto, Bureau of Commercial 
Fisheries, Seattle, Washington, sent us ten specimens of Sgualus 
acanthias taken from off the coast of Washington. W. I. Follett of 
the California Academy of Sciences, San Francisco, generously allowed 
us to use his and J. D. Hopkirk's vertebral counts of Cetorhinus maxi- 
mus and Carcharodon carcharias. These counts were made on verte- 
bral columns in the CAS collections. We are grateful to Sheldon 
Applegate, formerly of Duke University, for providing us with counts 
from New Jersey and Delaware specimens of Carcharias taurus, and 
to Elvira Siccardi, Faculty of Natural Sciences of Buenos Aires, 
Argentina, for counts on Isurus oxyrinchus, Carcharodon carcharias, 
and Lamna nasu^. 

Our identifications of the species of Sphyrna are the result of un- 
published studies by Dr. Carter R. Gilbert, to whom we extend our 
appreciation for allowing us to use his characterizations. 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 75 

Drs. Bruce B. Collette, Carl L. Hubbs, W. Ralph Taylor, and S. J. 
VVeitzman read the manuscript and made valuable suggestions for its 
improvement. 

Methods 

Sharks ranging in size from small embryos to specimens several feet 
long were used in the survey. Radiographs were made with a hard- 
ray machine and various industrial X-ray films. The finest-grained 
films gave the most desirable results, but generally grain size was a 
limiting factor only in making counts of the terminal vertebrae in the 
caudal fin. A few counts were made on skeletons. Each vertebral 
count was separated into two parts: 

1. Precaudal vertebral count (P) includes all complete centra 
anterior to the forward edge of the upper precaudal pit, or, in species 
where a pit is absent, all complete centra anterior to the origin of the 
upper lobe of the caudal fin. 

2. Caudal vertebral count (C) includes all centra posterior to the 
precaudal vertebrae. 

In order to demarcate clearly the precaudal from caudal centra on a 
radiograph, a pin was inserted at the forward edge of the upper pre- 
caudal pit, or at the upper caudal origin, so that its point touched the 
vertebral column. In some sharks, notably species of Brachaelums, 
Halaelurus, and Hemiscyllium, it is impossible to decide the point of 
origin of the dorsal lobe of the caudal fin; for these sharks, only total 
counts (T) are given. 

We do not know if our two methods of separating precaudal from 
caudal vertebrae produce homologus (hence comparative) counts in 
sharks with and without precaudal pits; however, the value of the 
methods lies in their usefulness and for the purposes of this study we 
consider the results homologous. Another possible way of subdivid- 
ing the vertebral column would be into monospondylous and diplo- 
spondylous centra.^ The transition from monospondyly (anterior) to 
diplospondyly (posterior) usually occurs above the pelvic fin, but there 
are notable exceptions (Etmopterus and a few species of CarcJiarhinus) 
wherein it is much further posterior. The transition usually is evi- 
denced by an abrupt reduction in centrum length (pi. 1a, b)^ but in 
several species, e.g., Alopias superciliosus (pi. Ic), Scoliodon laticaudus, 
Prionace glauca, the reduction is so slight as to be unnoticeable on a 
radiograph. 



2 Monospondyly=one centrum per myomere; diplospondyly=two centra per myomere. For detailed 
discussion, see Goodrich, 1930, pp. 26-27. 

• We have not made dissections to confirm the transition points from monospondyly to diplospondyly 
but accept the first notably shorter centrum above or behind the pelvic region as the first diplospondylous 
centrum. Confirmation of this view is found in some radiographs wherein it is possible to see the apertures 
for nerve roots Issuing: one pair for each centrum In the monospondylous region and one pair for two centra 
In the diplospondylous region. 



76 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Precaudal vertebral counts are subject to a maximum error of plus 
or minus two vertebrae. It is not always possible to decide accurately 
from a radiograph which is the first vertebra because, at the rear of 
the head, other structures, including the occipital condyles, mask the 
front of the vertebral column. Occasionally, owing to the angle at 
which our marking pin had entered, we had some doubt as to the last 
precaudal vertebra. Most of the counts given here were made by 
each of us working independently but using the same radiographs. 
The exceptions to these counts are the counts of the carcharhinid 
genera on which we are working individually. We never differed by 
more than one in our precaudal counts and we arbitrarily report the 
even count, appropriately compensating for the caudal count. 

At the tip of the caudal fin the last few centra of most shark species 
are frequently too small to give good resolution on radiographs. 
Sometimes there was a difference of as much as three or four vertebrae 
in our counts of the caudal section. In such counts one determina- 
tion or the other has been listed arbitrarily. Because of the range of 
variation in counts within a single species and because of the high 
number of vertebrae in most sharks, we believe that such an error 
is hardly significant. Generally, the precaudal count is more con- 
sistent than the caudal count. Caudal counts were made under 
magnification, usually with a low-power binocular microscope. 

The number of precaudal vertebrae is established early in embryos, 
but the last caudal vertebrae usually are not fully formed until late 
in embryonic life; hence, only late-stage embryos provide caudal 
counts comparable to those of adults. In those embryos in which 
we found calcification incomplete, we give a minimal count for the 
caudal vertebrae (!>). We doubt that more than 25 additional 
vertebrae would have been formed in any of the embryos we examined, 
and we believe that in most of the embryos the number of yet unformed 
vertebrae would be much fewer. 

Complete calcification of the column is recognizable on radio- 
graphs by the posterior extent reached by the column (to within a 
short distance of the tip of the tail) or by uniform density of the 
images of the last several centra. 

We counted apparent fusions between centra (and also apparent 
monospondylous intrusions in regions of diplospondyly) as single 
centra, but we recorded separately the recognizable centra when the 
fusion was obviously incomplete. 

In most sharks the monospondylous centra gradually increase in 
length from the cranium posteriorly, so that the centra are usually 
longest at, or near, the region where diplospondyly begins. The 
difference in the length of anterior and posterior monospondylous 
centra may be small or great. The difference in length between the 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 77 

posterior monospondylous centra and the anterior diplospondylous 
centra is similarly variable though reasonably constant for any one 
species. 

Reporting these relative centrum lengths is obviously of value but 
offers difficulties. Because many of the specimens we used were 
curled or distorted, radiogi-aphs of them show oblique images of 
some centra. The degree of obliqueness may vary from one region 
of the vertebral column to another; we, therefore, see the likelihood 
of introducing considerable error by giving dimensions of centra that 
are not adjacent to, or at least very close to, each other on the radio- 
graph. For this reason, we do not report the length of anterior 
relative to posterior monospondylous centra. To compare the length 
of a posterior monospondylous centrum with that of an anterior 
diplospondylous centrum (thereby indicating the relative prominence 
of the transition point between monospondyly and diplospondyly), 
we measured the length of the penultimate monospondylous centrum 
and the first diplospondylous centrum and then obtained a ratio, 
listed as "A" in the results and tables, by dividing the length of the 
former by the length of the latter and multiplying by 100. We 
chose the penultimate monospondylous centrum because it is usually 
more representative of the posterior monospondylous centra (the 
ultimate monospondylous centrum is often reduced in length even 
though noticeably longer than the first diplospondylous centrum). 
The first diplospondylous centrum was chosen because there is often 
a regular alternation of centrum length in the diplospondylous region. 
In deciding the transition point from monospondyly to diplospondyly 
when differences in length of the centra bordered on being imper- 
ceptible, we may be in error occasionally though we think that any 
error thus introduced is slight. 

To give an indication of centrum shape, we have divided the length 
of the penultimate monospondylous centrum by its diameter (appear- 
ing as height or width on the radiograph) and have multipled this 
ratio by 100 to yield the values listed as "B" in the results and tables. 

Results 

Our data on shark vertebrae are reported in table 1. Table 2 
condenses these data so that the subordinal, familial, and generic 
limits, but not those of the individual species, can be more easily 
visualized. 

We have representative samplings of six of the seven suborders of 
sharks and a partial sampling of the seventh. None of these sub- 
orders can be distinguished on its vertebral counts alone. In terms 
of total number of vertebrae we found a range of from 60 to about 419. 
The lowest total count is for Squaliolus laticaudus and the highest for 



78 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Alopias vulpinus. The species with the lowest precaudal count (44) 
is Squaliolus laticaudus and the one with the highest (149) is Prionace 
glauca. The lowest count (13) of caudal vertebrae is for Eupro- 
tomicrus bispinatus, Squaliolus laticaudus, and Squaliolus sarmenti, 
and the highest 0298) is for Alopias vulpinus. 

Compared with the other suborders, the Squaloidea have low 
vertebral counts. In all recent systems of classification the Squa- 
loidea are regarded as relatively advanced, so there is some justifica- 
tion for regarding theii- low counts as an advanced feature. On this 
basis, within the Squaloidea we could consider the generally lower 
counts of the dalatiids as indicating that they are more advanced 
than the squalids. The Notidanoidea and the Heterodontoidea, 
which, on other features, have some claim to being representative of 
the ancestral or primitive sharks, do not have more vertebrae than 
do many of the so-called advanced sharks in the Galeoidea. This 
may merely indicate that median numbers are most primitive, with 
both decrease and increase as specialized conditions. 

Only one family, the Alopiidae, with one genus, Alopias, is recog- 
nizable on its vertebral numbers alone. The distinctiveness of 
Alopias is due to the high number of vertebrae in its tail, a feature 
that might be expected from the extreme length of its tail compared 
to other sharks. The diagnostic value of vertebral numbers increases 
in the lower taxa. Thus, in the families containing several genera, 
a few genera {Lamna in the Lamnidae, Brachaelurus and Stegostoma 
in the Orectolobidae, Conoporoderma and Atelomycterus in the Scyli- 
orhinidae, Prionace in the Carcharhinidae, and perhaps others) are 
clearly recognizable on precaudal, caudal, or total count. Of the 
genera containing two or more species, almost half include at least 
one species distinguishable on vertebral counts. Because of our 
incomplete coverage, we believe it premature to list these species here. 

The suborders Squaloidea, Pristiophoroidea, Squatinoidea, and 
Heterodontoidea are consistent in having fewer caudal than precaudal 
vertebrae. In the suborder Galeoidea, the families Scyliorhinidae, 
Triakidae, and most of the Orectolobidae show the same trend. It 
may be of significance that the sharks with fewer precaudal than 
caudal vertebrae are for the most part small, and usually bottom- 
dwelling. In Carcharhinus (Garrick, in ms.), the large species tend 
to have fewer caudal than precaudal vertebrae and the small species 
have more caudal than precaudal vertebrae. 

In our studies, particularly of the genera Carcharhinus, Scoliodon, 
Loxodon, and Bhizoprionodon (Garrick, in ms.; Springer, 1964), we 
have noted no sexual dimorphism in vertebral numbers; however, 
Punnett (1904), who dissected 567 specimens of Etmopterus spinax 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 79 

from Norway, reported a statistically significant difference between 
males and females in numbers of monospondylous ("whole vertebrae") 
and diplospondyloiis ("half vertebrae") centra and in total number of 
segments (counting each whole vertebra or each two half vertebrae 
as one segment). Females had the higher average in each of these 
three categories but in none did the average difference exceed half a 
unit. Using Punnett's figures (for adults and embryos) we have 
plotted frequency distributions of total vertebrae (each whole or half 
vertebra of Punnett equals one vertebra in our study) and averaged 
them: 

Total vertebrae 81 82 83 84 85 86 87 88 89 90 91 Average 
Males 4 14 41 51 63 39 19 9 3 2 - 84. 73 

Females 3 12 39 71 84 55 33 20 3 - 2 85.02 

A t test of these data gives the value 2.06, which is not significant 
at the 95 percent level; however, a t test of the number of mono- 
spondylous vertebrae of adults gives a value which is significant 
above the 99.1 percent level. Punnett did not allow for possible 
year-class differences and it is not possible to tell from his data if there 
is bias. A study of sexual differences in vertebral counts based on 
embryos of a single year class would offer a solution to this problem. 

If the nature of sexual dimorphism in vertebral counts of sharks 
is generally of the magnitude shown by E. spinax, it is not surprising 
that our counts, based on small numbers of specimens, do not indicate it. 

mr A 1 / length penultimate monospondylous centrum ,^^„\ ti 

The A values ( — -, — ^,, a . ,■ , rh — 1 X100)> like 

V length first diplospondylous centrum / 

the vertebral numbers, vary widely within families and genera 

but within the species are relatively constant. The A values are 

smallest in the Lamnidae, Alopiidae, Triakidae, Carcharhinidae, and 

Sphyrnidae. The A values, up to about 300, are greatest in the 

Carcharhinidae, a family with great variation in this proportion. 

Families containing some species with A values of about 200 include 

the Scapanohynchidae, Triakidae, Sphyrnidae, Squalidae and Dala- 

tiidae. 

rpi Tj 1 /^r, / length penultimate monospondy lous centrum .yir,n\ 
JL ne jj values i -j-. r 7 tt'. — 7 t — ^ — — xioo 1 

\ diameter of penultimate monospondylous centrum / 

are in general as inconsistent within the families and genera as are the 
A values. The range of the B values is from about 25 {Alopias spp.) 
to 200 (Isistius hrasiliensis) . 

If a B value of 75 or less is arbitrarily considered low and one of 
125 or more is considered high, 13 of the 18 large sharks (attaining 
a size of over six feet) for which we have data are indicated as having 
low B values. Small species may have either high or low B values, 
but of the 30 species of sharks having high B values, only one, the 



80 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

unique Scapanorhynchus owstoni, attains a size greater than six feet. 
These facts seem to indicate that B values are functionally significant 
in terms of total length attained by the species. 

Remarks Concerning Certain Species 

In Hexanchus and Notorhynchus the vertebrae are not sufficiently 
calcified to produce an image on a radiogi'aph. In Heptranchias , the 
precaudal vertebrae are poorly calcified and there is considerable 
spacing between the images on the radiograph, which probably 
indicates that only the center or the ends of each centrum are registered. 
The caudal vertebrae are well calcified but terminate abruptly at about 
the level of the subterminal caudal notch in each specimen, a condition 
not noted in any other species we examined. 

In Chlamydoselachus anguineus only some of the vertebrae are 
calcified; hence, total counts could not be made. In one specimen the 
anterior 16 and posterior 3 precaudal centra and a few anterior caudal 
centra were apparent; in the other specimen radiographed, the anterior 
18 and posterior 9 precaudal centra and the anterior 20 caudal centra 
were apparent Goodey (1910) indicated the number of vertebrae in a 
specimen he dissected to be 112, on the basis of the number of neuro- 
meres, which were determined by counting the ventral-root foramina 
of the spinal nerves perforating the basidorsals. We believe that 
counting in this manner yields a figure which is much too low, for it 
does not take into account the diplospondylous condition. It is also 
obvious from Goodey's plates that no nerve roots issue from large 
segments of the vertebral column. 

Aasen (1961) reported vertebral counts of 60 specimens of Lamna 

nasus from the western North Atlantic, as follows: 

Total vertebrae 150 151 152 153 154 155 156 157 158 159 160 161 
Specimens 115698978141 

He noted that the number of precaudal vertebrae ranged from 83 to 89. 

One specimen of Paragaleus gruveli (USNM 196158) examined 
(pi. 1a) had two separated groups of elongated (monospondylous) 
centra in the precaudal region. If each of these elongated centra 
were counted as two, the number of precaudal centra would be 77 and 
the total count 147, which would be more in accord with the other 
specimen examined from the same locality. 

Specimens of Sphyrna tihuro from Florida have much lower vertebral 
counts than specimens from other localities. Counts from large num- 
bers of specimens throughout the Atlantic and Pacific range of this 
species should be made to determine the significance of the counts 
shown by the Florida specimens. 



PROC. U.S NAT. MUS. VOL 115 



SPRINGER AND GARRICK— PLATE 1 




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VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 81 

Squalus acanlhias is considered a cii-cumglobally distributed species 
in the temperate waters of both the northern and southern hemispheres 
although it is reported infrequently from subtropical areas. Our 
evidence (table 1) indicates that at least in the northern hemisphere 
the Atlantic population (with 79-85 precaudal vertebrae in 7 speci- 
mens) and the Pacific population (with 68-76 precaudal vertebrae in 
21 specimens) have differentiated. Six specimens from the southern 
hemisphere have 75-78 precaudal vertebrae and thus are intermediate 
between the northern hemisphere populations. A large-scale study 
of vertebral numbers in this species might yield interesting informa- 
tion on speciation. 

On the basis of the counts obtained for Centrophorus uyato, there 
is evidence that differences occur in the populations from the Gulf of 
Mexico and the eastern Atlantic. More counts will be necessary to 
verify this indication. 

The genus Etmopterus is d stinct from all other shark genera ex- 
amined in consistently having the last monospondylous centrum at 
a point behind the tip of the pelvic fin (pi. 1b), usually under the 
anterior portion of the second dorsal fin. In all other sharks we have 
investigated, except a few species of Carcharhinus (Garrick, in ms.), 
the last monospondylous vertebra occurs over the pelvic fin, usually 
over the pelvic base. 

Punnett (1904) reported the total vertebral counts of 163 free-living 
females of Etmopterus spinax. Twenty-five were gravid and were 
reported with the counts of theu* respective embryos (see our table 3). 
Examination of Punnett's data shows that the average number of 
vertebrae in a litter exceeds the maternal count in 10 of the 13 mothers 
with 82-85 vertebrae but is less than the maternal count in the 12 
mothers with 86-88 vertebrae. Differences in counts from a single 
litter range from two to seven vertebrae. We have found a similar 
variation in the range of vertebral counts in carcharhinid sharks in 
the few cases in which we have mothers with embryos. 

Only late-stage embryos of Echinorhinus cookei and E. brucus were 
radiographed; the radiographs show no evidence of calcification of 
the centra. 



727-653—64- 



82 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Literature Cited 

Aasen, Olav 

1961. Some observations on the biology of the porbeagle shark {Lamna 
nasus L.). Internat. Counc. Expl. Sea, Near Northern Seas Com- 
mitt., no. 109, pp. 1-7. [Mimeographed.] 
Bailey, Reeve M., and Gosline, William H. 

1955. Variation and systematic significance of vertebral counts in the 
American fishes of the family Percidae. Univ. Michigan Mus. 
Zool. Misc. Publ., no. 93, pp. 1-44. 

GOODEY, T. 

1910. A contribution to the skeletal anatomy of the frilled shark, 
Chlamydoselachus anguineus Gar. Proc. Zool. Soc. London, pp. 
540-570. 
Goodrich, E. S. 

1930. Studies on the structure and development of vertebrates, 837 pp. 
GtJNTHER, Albert 

1870. Catalogue of the fishes in the British Museum, vol. 8, xxv + 549 pp. 
HuBBs, Carl L., and McHugh, J. L. 

1951. Relationships of the pelagic shark Eu-protomicrus bispinatus, with 
description of a specimen from off California. Proc. California 
Acad. Sci., vol. 27, no. 6, pp. 159-176. 

ISHIYAMA, ReIZO 

1958. Studies on the rajid fishes (Rajidae) found in the waters around 
Japan. Journ. Shimonoseki College Fish., vol. 7, nos. 2 and 3, 
pp. 193-394. 
Jaquet, M. 

1905. Description de quelques parties du squelette du Pseudotriacis microdon 
Capello. Bull. Mus. Ocean. Monaco, vol. 36, pp. 1-28. 
Punnett, R, C. 

1904. Merism and sex in "Spinax niger." Biometrika, vol. 3, no. 4, pp. 
313-362. 
Schmidt, J. 

1917. Zoarces viviparus L. and local races of the same. Compt.-Rend. 
Trav. Labor. Carlsberg, vol. 13, no. 3, pp. 279-396. 
Springer, Victor G. 

1964. A revision of the carcharhinid shark genera Scoliodon, Loxodon, and 
Rhizoprionodon. Proc. U. S. Nat. Mus., vol. 115, no. 3493, 
pp. 559-632. 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 



83 



Table 1. — Vertebral characters in sharks 

P = precaudal vertebrae. C = caudal vertebrae. T = total vertebrae. A = 
length of penultimate monospondylous centrum divided by length of first 
diplospondylous centrum X 100. B = length of penultimate monospondylous 
centrum divided by its width X 100. 



Family, genus, species, locality 


P 


C 


T 


A 


B 


HEXANCHIDAE 












Hexanchus 












H. griseum (Bonnaterre) (see discussion) 


unc£ 


Icificd 








MCSN 34580, Italy 












USNM 186120, Florida 












Notorhynchus 












N. maculatus Ayres (see discussion) 


uncalcifled 








USNM 27071, California 












USNM 61234, California 












USNM 38301, Peru 












USNM 87681, Uruguay 












Heptranchias 












H.perlo (Bonnaterre) (see discussion) 












MCSN 37507, Ligurian Sea, Mediterranean 


ca. 85 


45 


ca. 130 






USNM 164167, south of Louisiana 


90 


66 


146 








89 


52 


141 






USNM 151972, ofl North Carolina 


90 


61 


151 






CHLAMYDOSELACHIDAE 












Chlamydoselachut 












C. anguineus Garman (see discussion) 






>112 


(Goodey, 


1910) 


USNM 48530, Japan 












USNM 161522, Japan 












HETERODONTIDAE 












Heterodontus 












H.francisci (Girard) 












USNM 34778, California 


71 


41 


112 


160 


95 


H. japonicus Dumerll 












USNM 71764, Japan 


72 


38 


110 


136 


94 


UMMZ 179075, Japan 


74 


42 


116 






H. galeatus (Giinther) 












USNM 59876, New South Wales, Australia 


71 


35 


106 


140 


92 


H. zebra (Gray) 












UMMZ 179077, locality unknown 




41 




133 


89 


CARCHARIIDAE 












Carcharias 












C. tauTUg Rafinesque 












USNM 143423, Louisiana, embryo 


84 


>70 


>154 






Sandy Hook Bay, New Jersey (from Sheldon 


82 


81 


163 






Applegate) 


80 


83 
84 
80 


163 








86 


166 








84 


79 


163 








84 


81 


165 








84 


- 


- 






1 mile ofl Lewes, Delaware (from Sheldon 


85 


71 


156 






Applegate) 


85 


85 


170 








83 


81 


164 








87 


81 


167 






8CAPAN0RHYNCHIDAE 












Scapanorhynchus 












S. owstoni (Jordan) 












MCZ 1279, Japan 


53 


69 


122 


189 


172 



84 



PROCEEDINGS OF THE NATIONAL MUSEUM 
Table 1. — Vertebral characters in sharks — Continued 



Family, genus, species, locality 



LAMNIDAE 

Isurm 
I. oxyrinchus Raflnesque 
MCZ 37994, Bahamas, embryo 
New York-New Jersey (from J. Q. Casey) 



Puerto Quequfo, Argentina (from E. Slccardl) 

/. glaucus 
UMMZ 177116, Japan 
SIO 50-240, California 

I. species (Qarrlck, In ms.) 
USNM 197427, Indo-Paclfic 
USNM 197429, Indo-Pacific 
Lamna 

L. nams (Bomiaterre) 
BMNH 1961.11.2.1, English Channel 
Puerto Quequ6n, Argentina (from E. Slccardl) 
Locality unspecified (after Gilnther, 1870) 
Western North Atlantic (after Aasen, 1961) 



Cnrcharodon 
C. carcharias (Linnaeus) 
CAS 26245, California 
CAS 26361, California 
CAS 26363, California 
CAS 26366, California 
CAS 26367, California 
CAS 26678, California 
CAS 26680, California 
CAS 26376, California 
CAS 26378, California 
CAS 26694, California 
CAS 26695, California 
CAS 26781, California 
CAS 26793, California 
CAS 27013, California 
CAS 27014. California 
CAS 27015, California 
Puerto Quequta, Argentina (from E. Slccardl) 

CETORHINIDAE 

Cetorhinus 
C. maximus (Qunnerus) 
CAS 25873, California 
USNM (uncat.), off British Columbia 
ALOPIDAE 
Alopias 
A. superciliosus (Lowe) 
UMML 8861, Florida, two slbllngcmbryos 

MCZ 36155, Cuba, embryo 
New York (from J. Q. Casey) 
USNM 197700, California 
A. vulpinus (Bonnaterre) 
CNHM 52100, Formosa 
SIO 52-19, QalSpagos, two sibling embryos 



110 


80 


190 


ca. 100 


no 


81 


191 




no 


80 


190 




109 


- 


- 




no 


86 


196 




108 


79 


187 


127 62 


112 


80 


192 


120 53 


112 


83 


195 


128 63 


111 


86 


197 


130 67 


85 


68 


153 


ca. 100 


91 


71 


162 




- 


- 


155 




-89 




150-161 


(based on 60 speci- 
mens; see text 
remarks) 


_ 


_ 


178 




104 


68 


172 




106 


68 


173 




104 


74 


178 




107 


ca. 76 


ca.l83 




107 


77 


184 




105 


ca.78 


ca. 183 




106 


73 


179 




106 


71 


177 




104 


77 


181 




104 


77 


181 




108 


73 


181 




103 


73 


176 




107 


74 


181 




105 


80 


185 




104 


76 


180 




104 


83 


187 




106 


79 


184 





50 



60 



110 
110 



102 


180 


282 






102 


187 


289 






102 


181 


283 ca. 


100 




102 


193±3 


295±3 






100 


204 


304 






121 


>298 


>419 






119 


243 


362 


136 


27 


119 


240 


359 


127 


26 



VERTEBRAL NUMBERS EN SHARKS — SPRINGER AND GARRICK 
Table 1. — Vertel)ral characters in sharks — (iJontinued 



85 



Family, genus, species, locality 



OEECTOLOBIDAE 
Oreetolobiu 
0. maculatus (Bonnaterre) 
USNM 176696, near Brisbane, Australia 
Stegoitoma 
S. tigrinum (Bonnaterre) 

RNH skeleton no. 429, Java 
S.fasciatum (Hermann) 
USNM 138547, Macassar market, Celebes 
USNM 138548, Macassar market, Celebes 
ChUoseyUium 
C. indkum (Qmelln) 

UMMZ 179022, Japan 
C. grUeum Miiller and Henle 

UMMZ 179034, Java 
C. species 

USNM 6449, Hong Kong 
C. species 
USNM 148107, Persian Qui/ 
USNM 148108, Persian Gulf 
EucTOssorhinui 
E. dasypogon (Bleeker) 
BMNH 1867.11.28.209, Aru Islands, syntype 
of Crossorhinus dasypogon Bleeker 
Qinglymostoma 
O. cirratum (Bonnaterre) 
USNM 196159, Senegal, sibling embryos 

USNM 37741, Jamaica (sibling embryos?) 

USNM 181312, Sonora, Mexico 
Hemiscyllium 
H. ocellatum (Bonnaterre) 
USNM 176863, Great Barrier Beef, 
Queensland, Australia 
H. sptciet 
USNM 123025, no locality 
BTOchaelUTUs 

B. waddi (Bloch and Schneider) 
USNM 197619, Sydney, Australia 

RHINCODONTIDAE 
Rhincodon 
R. typm Smith 
TGFC, off Texas, embryo 
SCYLIORHINIDAE 
Cephaloscyllium 

C. uter (Jordan and Gilbert) 
USNM 196142, California 

Oaleus 
Q. arae (Nichols) 
USNM 185602, oft Nicaragua (Atlantic) 

USNM 159233, ofl Florida 
USNM 168101, southwest of Grand Bahama 
Island 
HapJobkpharus 
H. edwardsi (Voigt) 
MCZ 1028, Cape St. Blaize, South Africa 



100 



100 



81 



60 



145 



62 



36 



166 



226 



162 



193 



>72 >153 



109 



133 



96 


122 


217 


160 


66 


101 


121 


222 


140 


68 


114 


61 


176 


143 


62 






170 


118 


72 


111 


01 


172 


116 


71 


104 


66 


169 


117 


74 


107 


60 


167 


117 


80 



97 


77 


174 


126 


80 


97 


73 


170 


118 


74 


98 


73 


171 


116 


71 


95 


75 


170 


121 


74 


92 


83 


175 


126 


66 






191 


120 


71 






190 


160 


76 



141 



75 


60 


135 


168 


143 


76 


>51 


>127 


173 


124 


74 


55 


129 


141 


124 


81 


69 


140 


147 


133 



43 



137 



107 



86 PROCEEDINGS OF THE NATIONAL MUSEUM 

Table 1. — Vertebral characters in sharks — Continued 



Family, genus, species, locality 



O 



SCYLIORHINIDAE-Continued 
Halaelurus 
H. vincenti (Zletz) 
UBC 56-406, Adelaide, South Australia 
ParmatUTUs 
P. xaniurus (Gilbert) 
MCZ 1002, off California 
ApristUTUs 
A, herklotsi (Fowler) 
USNM 93134, PhUippines, holotype of Pentan- 
chus herklotsi Fowler 
A. verweyi (Fowler) 
USNM 93135, Borneo, holotype of Pentanchus 
verweyi Fowler 
Scyliorhlnus 
S. torazame (Tanaka) 

UMMZ 179029, Fusan Market, Korea 
S. garmani (Fowler) 
USNM 43749, East Indies, hoMypc ol Halaelurus 
garmani Fowler 
Pentanchus 
P. profundicolus Fowler 
USNM 70260, Mindanao Sea, holotype (several 
centra missing, counts estimated) 
Conoporoderma 
C. marleyi (Fowler) 
ANSP 53427, Natal, South Africa, holotype of 
Poroderma marleyi Fowler 
Atelomycterug 
A. marmoratus (Bennett) 
UMMZ 179603, Java 
Undescrlbed genus and species (S. Springer, In ms.) 
USNM 185557, Caribbean Sea 
PSEUDOTRIAKIDAE 
Pseudotriakis 
P. microdon Capello 
Cape Verde Is. (from Jaquet, 1905) 
TRIAKIDAE 
ScyUiogaleus 
S. queketti Boulenger 
BMNH 1903.2.6.21, off Natal, South Africa, 
holotype 
Triakis 
T. semifasciata Olrard 

UMMZ 61065, California 
T. scyllia Miiller and Henle 
UMMZ 179099, Fusan Bay, Korea 

T. venustum (Tanaka) 

UMMZ 179065, Okinawa, Japan 
T. henlei (Gill) 

UMMZ 61051, California 

SU 34283, California 
Eridacnia 
E. radcliffei Smith 

USNM 74604, Philippines, holotype 



130 133 



70 60 130 145 



69 >50 >119 155 



70 46 116 139 



79 



110 
105 



84 



54 



28 



133 



104 



164 
145 



ca. 186 



>62 >140 



52 



136 



135 



172 



125 



114 



142 



114 



133 



74 


38 


112 


167 


105 


92 


40 


132 


162 


105 



113 



100 



125 



103 



117 



93 


60 


153 


200 


93 


93 


60 


153 


200 


95 


107 


61 


168 


144 


100 


109 


61 


160 


117 


83 


103 


49 


162 


112 


88 



47 



124 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 
Table 1. — Vertebral characters in sharks — Continued 



87 



Family, genus, species, locality 



TRIAKIDAE— Continued 
Mwitelwi 
M. manazo Bleeker 

UMMZ 178990, Japan 

UMMZ 178993, Japan 
M. kanekonis (Tanaka) 

UMMZ 179097, locality unknown 
M. canis (Mitchill) 

USNM 73091, New Jersey 
M. canis or M. schmitti Springer 

USNM 86724, Uruguay 

M. noTTtsi Springer 
USNM 116444, Florida 

M. higmani Springer and Lowe 
USNM 156930, oflE Siuinam, holotype 
USNM uncataloged, off British Oulana 
USNM uncataloged, off Surinam 



Triaenodon 
T. obeaus (Ruppell) 
ANSP 71738, Pearl and Hermes Reef, embryo 
Leptocharias 
L. smithi M tiller and Henle 
USNM 164435, Liberia 
MCZ 39691, off Banana, West Africa 
CARCHARHINIDAE 
CoTcharhinus 
C. sorrah (Valenciennes) 

9 specimens from China, Thailand, and Red Sea 
C. limbatiis (Valenciennes) 
5 specimens from Florida, Brazil, Virgin Islands, 
and Liberia 
C. Jalciformit (Bibron) 
14 specimens from western north Atlantic and 
eastern North Pacific Oceans 
C. leucas (Valenciennes) 
7 specimens from Florida, Lake Nicaragua, 
Guatemala, Tampico (Mexico), and Panama 
C. melanopierus (Quoy and Qaimard) 
14 specimens from GUbert, Caroline, and Philip- 
pine Islands and Thailand and Red Sea 
Aprimodon 
A. isodon (Muller and Henle) 
USNM 118457, Texas 
Negaprton 
N. breviTostris (Poey) 
CNHM 32743, Florida 
Hypoprion 
H. mactoti Miiller and Henle 
ISZZ 5799, Hong Kong 
SU 12988, Hong Kong 
SU 14488, Burma 
H. signatus Poey 
USNM 133827. Florida, embryo 



90 


52 


142 


136 


120 


00 


51 


141 


123 


112 


81 


49 


130 


153 


104 


90 


56 


146 


137 


111 


89 


48 


137 


169 


122 


90 


46 


136 


144 


121 


93 


52 


145 


137 


100 


88 


49 


137 


132 


102 


89 


50 


139 


131 


100 


87 


49 


136 


132 


100 


90 


51 


141 


154 


105 


89 


50 


139 


142 


92 


91 


53 


144 






129 


85 


214 


ca. 100 




137 


76 


213 


112 


68 


136 


75 


211 


ca. 100 





66-73 85-93 153-166 142-167 

96-100 98-101 194-198 110-138 

98-105 98-110 199-215 112-123 

110-114 95-104 208-218 110-130 

115-122 86-92 202-214 U2-136 



117 



84 



165 



201 



206 



125 



88-107 
62-89 

68-78 

60-72 

64-74 

102 
76 



70 


84 


154 


222 


154 


68 


82 


150 


191 


150 


70 


82 


152 


162 


124 



104 



80 



184 



120 



67 



88 PROCEEDINGS OF THE NATIONAL MUSEUM 

Table 1. — Vertebral characters in sharks — Continued 



Family, genus, species, locality 


P 





T 


A 


B 


CARCHARHINIDAE-Contlnued 












Rhizoprimodon (see Springer, 1964) 












Subgenus Rhizoprionodon 












R. acutus (RiippeU) 












121 specimens from the eastern Atlantic and 


55-79 


64-83 


121-162 






Indo-Pacific; counts vary according to locality. 












R. terraenovae (Richardson) 












Counts based on 74 specimens from the western 


58-66 


67-81 


126-144 


161-174 


118-145 


Atlantic from Nova Scotia to Yucatan. A and 












B values based on 6 specimens. 












R. poTosus (Poey) 












Counts based on 58 specimens from the western 


66-75 


69-85 


136-159 


125-141 


108-114 


Atlantic from the Bahamas to Uruguay. A 












and B values based on 5 specimens. 












R. longuTio (Jordan and Gilbert) 












Counts based on 39 specimens from the eastern 


68-86 


73-85 


146-167 






Pacific. 












Subgenus Protozygaena 












R. lalandei (Valenciennes) 












Counts based on 45 specimens from the western 


79-90 


67-79 


153-168 


126-138 


96-103 


Atlantic from Panama to Brazil. A and B 












values based on two specimens. 












R. oUgolini Springer 












Counts based on 57 specimens from the Indo- 


84-91 


64-75 


161-162 


ca. 100 




Pacific. 












R. taylori (Ogllby) 












Counts based on 10 specimens from Australia. 


73-80 


62-70 


135-149 


ca. 100 




Loxodon (see Springer, 1964) 












L. macTorhinm Miiller and Henle 












Counts based on 20 specimens, vary with locality; 


77-106 


71-86 


148-191 


112-208 


84-110 


A and B values based on 4 specimens 












Scoliodon (see Springer, 1964) 












S. lalicaudus Miiller and Henle 












Counts based on 98 specimens; A value based on 


97-112 


50-62 


148-171 


ca. 100 




4 specimens 












Prionace 












P. glauca (Llimaeus) 












BMNH 1961.11.2.2. English Chaimel 


149 


98 


247 


ca. 100 




USNM 48317, Italy 


143 


96 


239 






USNM 125766, off northern (eastern?) United 


142 


101 


243 






States 












USNM 164621-23, Hawaii, sibling embryos 


146 


106 


252 


110 


38 




146 


102 


248 








145 


102 


247 






UMMZ (field no. H29-15), Japan 


147 


>90 


>237 






USNM 197687, Capetown, South Africa 


143 


101 


244 






Gakorhinus 












0. galeus (Linnaeus) 












UZMK 463, locality unknown 


83 


53 


136 






0. japonicu3 (Miiller and Henle) 












UMMZ 179061, Fukuoka market, Japan 


100 


64 


160 


114 


82 


Hemipristis 












H. elongalus (Klunzinger) 












GVF 2386-1, Gulf of ThaUand 


104 


86 


190 








104 


90 


194 


122 


80 




103 


89 


192 


132 


64 


Oaleocerdo 












0. cuvieri (Peron and LeSueur) 












GVF 2357-1, Gulf of Thailand 


105 


126 


231 


113 


55 


USNM 196624, Florida, sibling embryos 


106 


>H7 


>223 


133 


62 




108 


>114 


>222 







VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 
Table 1. — Vertebral characters in sharks — Continued 



89 



Family, genus, species, locality 


P 





T 


A 


B 


CARCHARHINIDAE— Continued 












NegogaleM 












N. macrostoma (Bleeker) 












BMNH 1867.11.28.197, Java, holotype of HemU 


74 


63 


137 


130 


141 


galeus macrostoma Bleeker 












N. microstoma (Bleeker) 












BMNH 1867.11.28.173, Java, holotypo of Heml- 


81 


69 


150 


148 


108 


galeus microstoma Bleeker 












UMMZ 179017, Java 


78 


65 


143 






Paragaleus 












P. gruvtli Budker 












USNM 196158, Senegal, embryo (see discussion) 


72 


70 


142 


180 


140 


USNM 1961G3, Senegal, embryo (see discussion) 


79 


70 


149 


192 


144 


P. pectoralis (Qarman) 












M CZ 847, o2 New England (7) , holotype of Heml- 


78 


70 


148 


174 


iig 


galeus pectoralis Qarman 












SPHYRNIDAE 












Sphyrna 












S. tiburo (Linnaeus) 












USNM 88677, Guayaquil, Equador 


81 


82 


163 


158 


106 


USNM 190591, Panama Bay, Panama 






165 


145 


111 


ANSP 86208, San Miguel Bay, Panama 


86 


87 


173 


125 


89 


ANSP 581, Rhode Island 


88 


82 


170 


127 


93 


USNM 125763, Appalaehlcola Bay, Florida 


72 


70 


142 


165 


132 


USNM 116888, Tortuga.s, Florida 


72 


71 


143 


153 


124 


USNM 104318, Recife, Brazil 


80 


78 


158 


143 


111 


8. media Springer 












USNM 196140, Qulf of Nlcoya, Costa Rica 


103 


92 


195 


136 


72 




100 


93 


193 


120 


69 




103 


89 


192 








102 


88 


190 


111 


80 




101 


92 


193 


113 


65 




102 


92 


194 


150 


72 


USNM 190593, Panama Bay, Panama 


102 


94 


196 


112 


63 




102 


91 


193 








101 


92 


193 


116 


73 




101 


91 


192 


105 


72 


S. mokarran (RQppell) 












USNM 29645, Mazatlan, Mexico 


98 


108 


206 


137 


69 


USNM 40026, New South Wales, Australia, 












embyro 


97 


108 


205 


122 


78 


S. lewini (Griffith) 












USNM 72476, Java, embyro 


92 


105 


196 


156 


91 


USNM 130604. Foochow, China 


02 


100 


192 


148 


83 


USNM 29999, Jamaica 


96 


108 


204 






USNM 25180, South Carolina 


89 


85 


174 


155 


99 


S. zygaena (Linnaeus) 












UMMZ 179078. Japan 


99 


103 


202 


121 


82 


USNM 119699, Virginia 


102 


104 


206 


ca. 100 




S. tudei (Valenciennes) 












USNM 195957, Mississippi 


107 


95 


202 


ca.100 




S. blochi (Cuvler) 












RNH skeleton no. 343, Java 


52 


64 


116 






CNHM 21836, North Borneo 


54 


70 


124 


178 


128 


USNM 195846, Bombay, India 


52 


66 


118 


160 


133 



90 PROCEEDINGS OF THE NATIONAL MUSEUM 

Table 1. — Vertebral characters in sharks — Continued 



Family, genus, species, locality 


P 


C 


T 


A 


B 


SQUALID AE 












Squalus 












S. acanthias 












USNM174P5, Norway 


83 


26 


109 


179 


86 




83 


26 


109 


191 


75 


CNHM 35495-7, Scotland 


81 


30 


111 


169 


93 




85 


30 


115 


233 


97 




84 


29 


113 


173 


104 


USNM 75665. Virginia 


85 


32 


117 


140 


104 


USNM 121955. Virginia 


79 


29 


108 


118 


114 


USNM 197692, Capetown, South Africa 


75 


27 


102 






USNM 104828, Alaska 


70 


29 


99 


135 


108 




73 


30 


103 


162 


96 


USNM 52852, Alaska 


71 


29 


100 


141 


108 


USNM 27305, Puget Sound, Washington 


70 


30 


100 


155 


124 


USNM 50989, Puget Sound, Washington 


69 


28 


97 


141 


110 


USNM 197796, Washington 


73 


30 


103 








70 


29 


89 








71 


26 


97 








72 


27 


99 








70 


30 


100 








75 


29 


104 








72 


29 


101 








73 


31 


104 








76 


30 


106 








72 


28 


100 






USNM 71835, Japan 


68 


28 


96 


181 


97 




69 


29 


98 


167 


100 




72 


28 


100 


135 


100 


CNHM 59847, Japan 


70 


28 


98 


130 


107 


UMMZ 179056, Korea 


69 


29 


98 


148 


107 




71 


28 


99 


172 


108 


USNM 164430, ChUe 


78 


31 


109 


175 


131 




77 


30 


107 


150 


126 


USNM 164431, Chile 


76 


26 


102 


170 


131 


USNM 77306, Straits of Magellan 


76 


29 


105 


148 


137 




76 


31 


106 


151 


160 


S. blainvillei (Risso) 












USNM 130611, Ningapo, Chekiang, China 


86 


28 


114 


128 


80 


UMMZ 179055, Japan 


89 


29 


118 


156 


80 


USNM 196545, Taiwan 


89 


27 


116 


156 


86 


CNHM 55618, Korea 


89 


27 


116 


166 


81 


USNM 197393, Hawaiian Islands 


84 


29 


113 


129 


117 


USNM 157864, Florida 


87 


28 


115 


152 


98 




85 


30 


115 


155 


95 


USNM 160831, Florida 


86 


29 


115 


140 


96 


USNM 157748, Florida 


86 


30 


116 


150 


81 


USNM 168478, Louisiana 


86 


31 


117 


143 


83 


USNM 197691, Capetown, South Africa 


82 


32 


117 






MCSN 34458, Italy 


80 


26 


106 


160 


100 


USNM 28473 (presumably Italy) 


78 


28 


106 


155 


102 




79 


29 


108 


165 


94 


S. megalops (Macleay) 












GVF HK95 south China Sea 


80 


25 


105 








78 


26 


104 








78 










USNM 191180, Taiwan 


81 


26 


107 


160 


100 




82 


26 


108 


136 


94 




81 


27 


108 


165 


93 


UMMZ 179033, Japan 


80 


26 


106 


136 


97 


UMMZ, 179053, Japan 


79 


27 


106 


144 


82 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 
Table 1. — Vertebral characters in sharks — CJontinued 



91 



Family, genus, species, locality 


P 


C 


T 


A 


B 


SQUALIDAE— Continued 












Sgualus— Continued 












S. cubensis Howell-Rivero 












USNM 157853, north of Cuba 


84 


30 


114 


150 


80 


USNM 157846, north of Cuba 


86 


30 


116 


141 


100 


USNM 196544, north of Cuba 


87 


29 


116 


144 


92 


USNM 164247, Dominican Republic 


82 


30 


112 






CentrophoTus 












C. uyato (Raflnesque) 












USNM 196160, Senegal 


90 


32 


122 


136 


95 


USNM 157862, south of Alabama 


82 


31 


113 








83 


30 


113 






C. squamosus (Bonnaterre) 












USNM 161517-18, Japan 


82 


24 


106 








83 


24 


107 


125 


78 


MCZ 40769, New Zealand 


86 


28 


114 


122 


100 


Centrosci/mnus 












C. owstoni Oarman 












UMMZ 142885, Japan 


74 


28 


102 


125 


69 


C. crepidateT Bocage and Capello 












USNM 196151, Senegal 


77 


29 


106 


137 


128 


MCZ 40768, New Zealand, embryo, uncalcified 












C. coelolepis Bocage and Capello 












USNM 38072, oft New Jersey 


78 


27 


105 


133 


80 


MCZ 37420, east of New York 


79 


26 


105 


126 


67 


MCZ 37424, east of New York 


79 


29 


108 


122 


65 


USNM 94522, Madeira 


75 


34 


109 


125 


118 


CentrosajUium, 












C. fabricii (Reinhardt) 












USNM 131383, off Newfoundland 


66 


29 


95 


162 


130 


USNM 38110, off New York 


67 


30 


97 


200 


122 


C. Titlen Jordan and Fowler 












UMMZ 179024, off Japan 


62 


28 


90 


141 


124 


USNM 161521, Japan 


61 


27 


88 


181 


106 




64 


27 


91 


161 


119 




64 


26 


90 


160 


121 


Deania 












D. elegans Springer 












USNM 159603, off North Carolina, holotype 


85 


34 


19 


133 


80 


D. calcea (Lowe) 












UMMZ 142884, Japan 


90 


36 


126 






Elmopleria 












E. polli Blgelow, Schroeder, and Springer 












USNM 163370, off Angola 


54 


24 


78 


144 


163 


E. spinax (Linnaeus) 












USNM 195848, Italy 


55 


29 


84 


130 


118 


Counts based on 567 specimens from Norway 






81-91 






(after Punnett, 1904; see our discussion) 












E. mens Bigelow, Schroeder, and Springer 












USNM 158406, Gulf of Campeche, Mexico 


56 


27 


83 


173 


173 


USNM 185600, Nicaragua (Atlantic) 


57 


24 


81 


144 


144 




67 


24 


81 


140 


140 




58 


26 


84 


150 


150 


E. schuUzi Bigelow, Schroeder, and Springer 












USNM 158144, south of Alabama 


56 


26 


82 


160 


120 




67 


25 


82 


153 


153 




66 


26 


82 


143 


133 




58 


24 


82 


147 


147 




56 


27 


83 


143 


165 



92 PROCEEDINGS OF THE NATIONAL MUSEUM 

Table 1. — Vertehral characters in sharks — Continued 



Family, genus, species, locality 



SQUALIDAE— Continued 

Etmopterm—Coutinned 

E. princeps CoUett 

USNM 163365, off New Jersey 
E. lucifer Jordan and Snyder 
USNM 51282, Japan 

USNM 161515, Japan 
E. buUisi Blgelow and Schroeder 
USNM 185597, northeast of Honduras 

USNM 185603, North Carolina 
E. baxUri Oarrick 

MOZ 40688, New Zealand 
E. puiiUus Lowe 

USNM 157835, off northwest Florida 
E. frontimaculatui Pletschmann 

USNM 196521, Japan 

UNMZ 179025, Japan 
OXYNOTIDAE 
Oxynotus 

0. centrina (Linnaeus) 
ANSP 575-76 Italy 

DALATIIDAE 

Dalatias 

D. licha (Bonnaterre) 
MCSN 34631, Italy 
MCZ 910, Italy 

USNM 157844, south of Alabama 
Scymnodalatias 
S. sherwoodi (Archey) 
CM no number, New Zealand, holotype of 
Scymnodon sherwoodi Archey 
EupTotomicTUi 

E. bispinatus (Quoy and Galmard) 
USNM 190031, near Midway Island 
USNM 164176, west of Johnston Island 
USNM 190032, south Pacific Ocean 

CAS 20431, off California (after Hubbs and 
McHugh, 1951) 
Heteroscymnoides 
H. marleyi Fowler 
ANSP 53046, South Africa, holotype 
MbHub 

1. brasilensis (Quoy and Galmard) 
USNM 164174, west of Christmas Island 
USNM 164175, northwest of Fanning Island 
USNM 190039, near Fanning Island 

USNM uncataloged, off Mississippi (Oregon 

2507) 
USNM uncataloged, off Mississippi (Oregon 

2945) 
I. species (Oarrick and S. Springer, in ms.) 
USNM uncataloged, oS Mississippi (Oregon 

3103) 



58 



58 



48 



52 



65 



23 



23 



18 



27 



81 



70 



92 



211 



143 



130 



160 



167 



136 



59 


26 


85 


154 


176 


62 


27 


89 


152 


173 


63 


27 


90 






60 


24 


84 


176 


131 


61 


24 


85 


178 


114 


60 


24 


84 


150 


160 


61 


27 


88 


141 


135 


64 


24 


88 


154 


180 


66 


22 


88 


127 


136 


61 











51 


27 


78 


140 


156 


51 


28 


79 


143 


152 


60 


28 


78 


153 


149 



152 



51 


13 


64 


182 


190 


47 


14 


61 


173 


161 


46 


15 


61 


130 


144 



14S 



61 


24 


85 


157 


147 


61 


25 


86 


150 


161 


60 


24 


84 


166 


200 


62 


20 


82 


172 


172 


61 


20 


81 


159 


189 


66 


23 


89 


162 


179 



180 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 
Table 1. — Vertebral characters in sharks — Continued 



93 



Family, genus, species, locality 



T 



DALATHDAE— Continued 
Sgualiolas 
S. sarmenii Noronja 

ANSP 2454, Madeira 
jS. lalicaudus Smith and Radellffe 
USNM 70259, Luzon, Philippines, holotype 
USNM 76679, Luzon, Philippines, paratype 
Somniosus 
S. Tostratus Capello 
IRSN 1399c, Mediterranean 
ECHINORHINIDAE 
Echinorhinus (see discussion) 
E. cookei Pletschmann 

USNM 179805, Hawaii, embryos 
E. brucus (Bormaterre) 
BMNH 1891.7.2.3.1, Nice, embryo 
PRISTIOPHORIDAE 
PristiophoTua 
P. ichroederi Springer and BuUls 

UMMZ 178848, ofl Cuba 
P. oweni Qiinther 

BMNH 1859.9.11.1, locality unknown, holotype 
P. japonictis Qiinther 
BMNH 1862.11.1.37, Japan, syntype 
BMNH no number, Japan, syntype 
UMMZ 176819, Japan 
P. nudipinnis Qiinther 
BMNH 1869.2.24.2, Tasmania, syntype 
Pliotrtma 
P. warreni Regan 
BMNH 1905.6.8.9, Natal, South Africa, holotype 
SQUATINIDAE 
Sguaiina 
S. japonica Bleeker 

UMMZ 179041, Nagasaki market, Japan 
<S. sguatina (Linnaeus) 
IRSN 1401B, Mediterranean 
RNH skeleton no. 492, Holland 
RNH skeleton no. 428, no locality 
iS. californica Ayres 
UBC 59-237, Cape San Lucas, Baja 
California, Mexico 



48 


13 


61 


139 


178 


47 


13 


60 


173 


173 


44 






150 


150 



100 


55 


155 


144 


100 


103 


48 


151 


150 


78 


109 


40 


149 


114 


83 


104 


63 


157 






108 


48 


166 


143 


75 


103 


50 


153 


117 


99 



106 



49 



155 



02 


34 


126 


133 


93 


28 


121 




96 








93 








108 


33 


141 


135 


108 


32 


140 


135 



83 



70 



94 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 2. — Summary of vertebral characters in suborders, families, and genera of 

sharks 

For explanation of symbols, see table 1 





P 


C 


T 




A 


B 


NOTIDANOIDEA 














Hexanchidae 














Hezanchus 






ancalcifled 








Noiorht/nchus 






unealcifled 








Hepiranchias 


85-90 


45-61 


ca. 130-161 








CnLAMYDOSELACHOIDEA 














Chlamydoselachidae 














Ctilamydoselachus 






>112 








IlETERODONTOIDEA 














Heteiodontidae 














Heterodonius 


71-74 


35-42 


100-116 




133-160 


89-95 


OALEOIDEA 


50-149 


28->298 


104->419 


ca 


100-328 


25-172 


Carchariidae 














Carcharias 


80-87 


71-85 


150-170 








Scapanorhynchidae 














Scapan orhyn dm s 


53 


69 


122 




189 


172 


Lamuldae 


84^112 


68-86 


160-197 


ca. 


100-130 


52-67 


ISUTUS 


108-112 


79-86 


187-197 




100-130 


62-67 


Lamna 


84-91 


68-71 


160-162 


ca 


. 100 




Carcharodon 


103-108 


68-83 


172-187 








Cetorhinidae 














Cetorhinus 


50 


60 


110 








Alopiidae 














Alopias 


100-121 


180->298 


282->419 


ca. 


100-136 


25-27 


Orectolobidae 


81-114 


50-145 


142-226 




115-150 


55-80 


Orectolobus 


106 


60 


166 




133 


78 


Stegostoma 


81-101 


121-145 


217-226 




140-150 


55-58 


Chiloscyllium 


104-114 


60-65 


167-175 




115-143 


62-80 


Eucrossorhinus 


100 


62 


162 








Ginglymostoma 


92-98 


73-83 


170-175 




116-125 


66-80 


Hemiscyllium 






190-192 




120-150 


71-75 


Brachaelurus 






142 




125 


68 


Rhineodontidae 














jRhincodon 


81 


>72 


>153 








Scyliorhinidae 


69-110 


28-60 


104-164 




133-173 


94-143 


Cephaloscylliu m 


74 


35 


109 




141 


94 


Oaleus 


74-81 


> 51-60 


>127-140 




141-173 


124-143 


Haploblephams 


94 


43 


137 




140 


107 


Halaelurus 






130 




133 


114 


ParmatuTUS 


70 


60 


130 




145 


115 


Apristurus 


69-70 


46->60 


116-M19 




139-155 


122-133 


Sctjliorhinus 


74-92 


38-40 


112-132 




162-167 


105 


Pentanchus 


79 


54 


133 




135 


113 


Conoporoderma 


76 


28 


104 




172 


100 


AtelomycUrus 


110 


54 


164 








Undescribed genus (S. Springer, in ms.) 


105 


40 


145 




125 


125 


Pseudotriakidae 














Pseudotriakis (from Jaquct, 1905) 






cal86 








Trlakidae 


77-137 


46-85 


124-214 


ca 


100-200 


68-122 


Scylliogaleus 


88 


>52 


>140 




114 


103 


Triakis 


84-109 


49-60 


136-160 




112-200 


83-117 


Eridacnis 


77 


47 


124 




125 


105 


Mustelus 


81-93 


46-56 


130-146 




123-169 


92-122 


Triaenodon 


129 


85 


214 


ca 


100 




Leptocharia 


136-137 


75-76 


211-213 


ca 


100-112 


68 



VERTEBRAL NUMBERS IN SHARKS — SPRINGER AND GARRICK 



95 



Table 2. — Summary of vetehral characters in suborders, families, and genera of 

sharks — Continued 

For explanation of symbols, see table 1 





P 


C 


T 


A 


B 


OALEOIDE A— Continued 












Carebarhlnidae 


54-149 


50-126 


110-252 


ca. 100-328 


38-154 


Carcharhinus (Qarrlck, in ms.)* 


54-125 


53-110 


110-235 


110-328 


60-107 


Aprionodon 


79 


86 


165 


206 


150 


Negaprion 


117 


84 


201 


125 


76 


Hypoprion 


70-104 


80-86 


152-184 


120-222 


67-154 


Rhizoprionodon 












subgenus Rhizoprionodon 


55-86 


64-85 


121-167 


125-174 


108-145 


subgenus Protozygaena 


73-91 


62-79 


135-168 


ca. 100-138 


96-103 


Loxodon 


77-106 


71-86 


148-191 


112-208 


84-119 


Scoliodon 


97-112 


50-62 


148-171 


ca. 100 




Prionace 


142-149 


>90-106 


>237-252 


110 


38 


Qalcorhinus 


83-106 


53-54 


136-100 


114 


82 


Ilemipristis 


103-104 


86-90 


190-194 


122-132 


64-80 


Galeocerdo 


105-108 


>114-126 


>222-231 


113-133 


65-62 


Negogaleus 


74-81 


63-69 


137-150 


139-148 


108-141 


Paragaleus 


7:^-79 


70 


142-149 


174-192 


119-144 


Sphymidae 












Sphyrna 


52-107 


64-108 


116-206 


ca. 100-178 


63-133 


SQUALOIDEA 


44-90 


13-36 


60-126 


118-233 


65-200 


Squalidae 


54-90 


22-36 


78-126 


118-233 


65-180 


Squalm 


68-89 


26-32 


96-118 


118-233 


75-160 


Centrophorus 


82-90 


24-32 


106-122 


122-136 


7^-100 


Centroscymnus 


74-79 


26-34 


102-109 


122-137 


65-128 


Ceniroscylliiim 


61-67 


26-30 


88-97 


141-200 


106-130 


Deania 


85-90 


34-36 


119-126 


133 


80 


Etmopterus 


54-66 


22-29 


78-91 


127-211 


114-180 


Oxj'notidae 












Oxynotus 


61-64 


30 


91-94 


143 


05 


Dalatlidae 


44-66 


13-28 


60-92 


130-182 


144-200 


Balatias 


50-51 


27-28 


78-79 


140-153 


149-156 


Scymnod alalia s 


58 


23 


81 


130 


152 


Euprotomicrtis 


46-51 


13-15 


61-64 


130-182 


144-190 


Heieroscymnoides 


52 


18 


70 


160 


145 


Jsistius 


60-06 


20-27 


81-92 


150-172 


147-200 


Squaliolus 


44-48 


13 


60-61 


139-173 


150-178 


Somniosus 


58 


20 


78 






Ecbinorhinidae 












Echinorhinus 


un calcified 








PRISTIOPHOIIOIDEA 












Pristiophoridae 


100-109 


40-55 


149-157 


114-150 


75-100 


PristiophoTus 


100-109 


40-55 


149-157 


114-150 


75-100 


Pliolrema 


106 


49 


155 


121 


83 


SQUATINOIDEA 












Squatinldae 












Squatina 


92-108 


28-34 


121-141 


133-135 


53-70 



•Includes data not in table 1. 



96 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 3. — Etmopterus spinax: total vertebral numbers in females with litters 
(based on data in Punnett, 1904) 



Total vertebral count 


Number of sibling 


Mother 


Average of sibling 
embryos 


Range of sibling 
embryos 


embryos 


82 


84.7 


83-87 


6 


82 


84.7 


83-87 


7 


83 


83.6 


81-86 


12 


83 


84.2 


82-86 


9 


84 


83.3 


82-85 


7 


84 


83.9 


83-85 


10 


84 


84.3 


83-86 


6 


84 


84.5 


82-87 


12 


84 


85.5 


84-88 


6 


85 


83.8 


82-85 


8 


85 


85.2 


84-88 


13 


85 


85.3 


85-87 


6 


85 


85.5 


85-86 


2 


86 


83.7 


82-87 


10 


86 


83.9 


82-B6 


9 


86 


84.5 


82-87 


11 


86 


84.7 


84-87 


9 


86 


84.8 


84-86 


9 


86 


84.9 


83-87 


8 


86 


85.2 


84-«7 


13 


86 


85.6 


85-87 


14 


87 


84.2 


83-«5 


8 


87 


86.8 


85-88 


12 


88 


86.0 


84-88 


8 


88 


87.2 


84-90 


9 



U.S. GOVERNMENT PRINTINC OFFICEll964 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION . WASHINGTON, B.C. 

Volume 116 1964 Number 3497 

NEOTROPICAL MICROLEPIDOPTERA, IV 

A NEW GENUS OF STENOMIDAE 

WITH DESCRIPTIONS OF FOUR NEW SPECIES 

(LEPIDOPTERA: GELECHIOIDEA) 



By W. Donald Duckworth 



This, the fourth in a series of papers ^ dealing with Neotropical 
moths, is the first of a planned series on the family Stenomidae, The 
aim of this research program is a reclassification of the family Steno- 
midae that more adequately reflects the natural relationships of the 
species. The present plan is to continue to expand research already 
in progress on the genera and to conduct detailed studies of the 
species. 

Since the genus Stenoma Zeller was proposed, it has become a 
"waste basket" for hundreds of species that could not be assigned 
readily to other genera on wing or palpal characters. Obviously this 
heterogeneity must be eliminated if the classification is to reflect 
natural relationships. In order to achieve this end, I find it necessary 
to regroup the species now assigned to the genus Stenoma according 
to their natural affinities. 

The new genus described here is composed of ten species formerly 

' Prepared with the aid of a National Science Foundation grant. Previous parts 
of this same series are: I and II, J. F. Gates Clarke, 1962, Proc. U.S. Nat. Mus., 
vol. 113, no. 3457, pp. 373-388; III, Clarke, 1964,ibid., vol. 115, no. 3480, pp. 61-84. 

97 

727-643 — 64 



98 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii8 

placed in Stenoma, of which one {S. curiata) is a synonym, together 
with four new species encountered during the course of this study. 
Unfortunately, as is the case for most species of Stenomidae, host 
plant records and other biological data are lacking; however, I hope 
that field explorations planned for the near future will correct this 
situation at least partially. 

This research was aided by a grant from the National Science 
Foundation. The drawings were made by Mr. Andre Pizzini and the 
photographs by Mr. Jack Scott, staff photographer. 

Genus Lethata, new genus 

Type-species: Stenoma trochalosticta Walsingham, 1913, in Godman 
and Salvin, Biologia Centrali-Americana, vol. 42 (Lepidoptera- 
Heterocera, vol. 4), p. 177. 

Head with spreading lateral tufts. Labial palpus long, recurved; 
second segment thickened with appressed scales; apical segment acute, 
shorter than second. Forewing with costa arched, apex rounded, 
termen obliquely rounded, tornus rounded; 12 veins, all separate: 
2 from before angle, 3 from angle, 4 approximate to 3, 7 to termen, 8 
to apex. Hindwing broader than forewing; 8 veins: 3 and 4 connate, 
6 and 7 connate. Abdomen of males with strong, eversible pair of 
hair-pencils contained in elongate cuplike pocket on fii*st sternum 
(Fig. 3a). 

Male genitalia: Uncus short, more or less recurved; tegumen some- 
what narrowed in the area of the uncus and gnathos; gnathos fused, 
setiferous; harpe simple, bifurcate setae present or absent; anellar 
lobes symmetrical or asymmetrical; aedeagus with or without cornuti. 

Female genitalia: Ostium bursae sclerotized; ductus birrsae sclero- 
tized to inception of ductus seminalis, membranous beyond; corpus 
bursae membranous, with two dentate signa connected by narrow, 
weakly sclerotized band. 

This genus is distinguished readily from all other genera in the 
famUy Stenomidae by the setiferous gnathos and the two dentate 
signa connected by a weakly sclerotized band in the corpus bursae. 

Although exact relationships are virtually impossible to ascertain 
at this time, the general structure of the genitalia in the genus Lethata 
indicates an affinity with Gonioterma Walsingham. 

One of the most striking features of the family Stenomidae is the 
presence of modified setae on the harpes of the male genitalia. In 
the genus Lethata, however, apparently this character is not consistent. 
Initially it was thought that modified setae did not occur in the genus 
at all, but examination at very high magnifications revealed the tips 
of the setae to be bifurcate in some species. A seta of this type has 



NEOTROPICAL MICROLEPIDOPTERA — DUCKWORTH 99 

been illustrated in figure 2j. Other species were found lacking this 
condition, and since it is possible that the tips of the setae might be 
broken during the preparation of slides of the genitalia, definite con- 
clusions concerning their presence or absence must await examination 
of additional material. 

Key to the Species of Lethata Based on the Genitalia 

1. Male 2 

Female 14 

2. Anellar lobes symmetrical 3 

Anellar lobes asymmetrical 13 

3. Vesica armed with coniuti 4 

Vesica without cornuti 12 

4. Vesica armed with small spiculate cornuti 5 

Vesica armed with one or more large, heavily sclerotized cornuti ... 7 

5. Harpe with saccular lobe bearing two long, heavily sclerotized setae. 

L. trochilosticta (Walsingham) 
Harpe not as above 6 

6. Anellar lobes approximately triangular with single, short spine at apex; 

aedeagus without spine, apex blunt L. fusca, new species 

Anellar lobes narrow with large, strongly recurved, hooklike spine at apex; 
aedeagus with posterior spine, apex sharply pointed. 

L. satyropa (Meyrick) 

7. Aedeagus with apical process; anellar lobes narrowing sharply at middle 

forming narrow, apically acute projections 8 

Aedeagus without apical process; anellar lobes not narrowing sharply at 
middle 9 

8. Aedeagus with apical process hooklike L. buscki, new species 

Aedeagus with apical process straight L. asthenopa (Meyrick) 

9. Aedeagus with vesica armed with more than six cornuti 10 

Aedeagus with vesica armed with less than six cornuti 11 

10. Aedeagus large, broad basally, vesica armed with ladder-like series of large to 

small cornuti extending from base to apex; anellar lobes of near uniform 

width throughout L. invigilans (Meyrick) 

Aedeagus small, narrow, vesica armed with two clusters of small cornuti at 
apex and single, large cornutus basally; anellar lobes approximately 
triangular in shape, pointed at apex . . . . L. maculata, new species 

11. Anellar lobes serrate beyond apical third; harpe broader basally. 

L. bovinella (Busck) 
Anellar lobes not serrate; harpe of approximately equal width throughout. 

L. leucothea (Busck) 

12. Aedeagus with two laterally directed processes at apex, one extending from 

each side, the larger one slightly recurved . . . L. glancopa (Meyrick) 
Aedeagus with large, heavily sclerotized process posteriorly. 

L. pyrenodes (Meyrick) 

13. One anellar lobe long, narrow, curved 90° laterad at middle and extending 

to base of opposite harpe; the other shorter, broader, sharply recurved; 
aedeagus with large, recurved spine at apex . . L. aromatica (Meyrick) 
One anellar lobe narrow, pointed at apex; the other broad with three large 
spines apically; aedeagus with apex on one side produced into long blade- 
like process L. ruba, new species 



100 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

14. Inception of ductus seminalis near ostium bursae; ductus bursae with 

corrugated, pouchlike area just before ostium . L. maculata, new species 

Inception of ductus seminalis near corpus bursae; ductus bursae not as 

above 15 

15. Ductus bursae with corrugated, ribbon-like sclerotized area from ostium to 

inception of ductus seminalis L. invigilans (Meyrick) 

Ductus bursae with patch of small spines near inception of ductus seminalis. 

L. bovinella (Busck) 

Lethata trochalosticta (Walsingham), new combination 

Figures \h-d, 4a 

Stenoma trochalosticta Walsingham, 1913, in Godman and Salvin, Biologia 
Centrali- Americana, vol. 42 (Lepidoptera-Heterocera, vol. 4), p. 177. 

Alar expanse 36 mm. 

Antenna brownish. Head brownish gray shaded with rose; second 
segment of labial palpus rose on outer side, whitish on inner side, 
apical segment whitish. Legs whitish shaded with rose. Thorax deep 
ochreous with brownish-yellow tuft of white-tipped scales medially. 
Forewing deep ochreous with costa rosy red to apical third; from 
middle of base a reddish-brown shade blends outward and downward 
with ground color; spot at end of cell composed of ring of reddish 
brown enclosing spot of rosy red with fuscous spot in its center; cilia 
dark gray basally, lighter beyond. Hindwing whitish ochreous slightly 
overcast with gray scales; cilia rosy. 

Male genitalia (slide WDD 2100, type): Uncus bluntly pointed 
apically, shghtly recurved; gnathos short, apex flattened dorsally; 
harpe broad basally, sacculus produced into lobe that bears two long, 
heavily sclerotized setae, apex rounded; anellar lobes symmetrical, 
slightly recurved and bearing long setae apically; aedeagus with 
ventral lip of apex extending beyond dorsal lip and acute at apex, 
vesica armed with numerous spiculate cornuti. 

Female genitalia: Unknown. 

Type: In the United States National Museum. 

Type locality : Chiriqui, Panama. 

Distribution: Panama: Chiriqui. Costarica: Sixaola River. 

This species may be distinguished from other species of Lethata by 
the conspicuous ocellate spot at the end of the cell in the forewing 
and the saccular lobe bearing two long, heavily sclerotized setae on 
the harpe of the male genitalia. 

Lethata aromatica (Meyrick), new combination 

Figures lk,l, 46 
Stenoma aromatica Meyrick, 1915, Exotic Microlepidoptera, vol. 1, p 449. 
Alar expanse 21-22 mm. 
Antenna brown. Head brown; labial palpus whitish ochreous. 



NEOTROPICAL MICROLEPIDOPTERA — DUCKWORTH 101 

Legs white shaded with brown . Thorax and tegula brownish ochreous, 
with scattered fuscous scales; patches of brown scales at inner angle 
and along dorsum; faint transverse line from dorsum to middle at 
apical third; cilia brown basally, lighter beyond. Hindwing whitish, 
dorsal half shaded with dark scales; cilia whitish. 

Male genitaha (slide WDD 3046) : Uncus pointed apically, shghtly 
recurved; gnathos long, finger-hke; harpe broad basally, tapering 
to pointed apex; anellar lobes asymmetrical; one long, narrow, 
curved 90° laterad at middle and extended to base of opposite harpe; 
the other shorter, broader, sharply recurved; aedeagus with large 
recurved spine at apex, vesica without cornuti. 

Female genitalia: Unknown. 

Type: In the British Museum (Natural History). 

Type locaUty: Sao Paulo, Brazil. 

Distribution: brazil: Sao Paulo; Parang. 

This species is known only from the male type in the British Mu- 
seum (Natural History) and one male specimen from Parana, Brazil, 
in the collection of the United States Natonal Museum. 

The species is easily recognized by the striking development of the 
anellar lobes in the male genitalia. 

Lethata asthenopa (Meyrick), new combination 

Stenoma asthenopa Meyrick, 1916, Exotic Microlepidoptera, vol. 1, p. 517. 

This species is known only from the male type in the British Mu- 
seum (Natural History), which has not been available for study; 
however, Clarke (1955, p. 212, pi. 106, figs. 2-2b) illustrates the wings 
and male genitalia. From these illustrations it is apparent that S. 
asthenopa belongs in the genus Lethata and is specifically distinct 
from the other species. Judging from the original description, S. 
asthenopa is closest to L. jusca, new species, in color and maculation. 
The color of the forewing of L. asthenopa is somewhat lighter brown 
than that of L. fusca. The genitalia afford a number of distinguish- 
ing characters, particularly the shape of the anellar lobes and the 
shape and armament of the aedeagus. 

Type: In the British Museum (Natural History). 

Type locality: Maroni River, French Guiana. 

Distribution: French guiana: Maroni River. 

Lethata bovinella (Busck), new combination 

FiGUBEs li,j, 3d, 4c 

Stenoma bovinella Busck, 1914, Proc. U.S. Nat. Mus., vol. 47, p. 50. 
Stenoma curiata Meyrick, 1929, Trans. Ent. Soc. London, vol. 76, p. 515. (New 
synonymy.) 

Alar expanse 19-20 mm. 



102 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

Antenna reddish brown basally, brown beyond. Head reddish 
brown, hghter between eyes; second segment of labial palpus whitish 
on inner side, brown on outer side, apical segment whitish. Legs 
whitish, shaded with brown. Thorax rosy ochreous dorsally, with 
central, posterior ridge of raised reddish brown scales. Forewings 
rosy ochreous with costa narrowly brick red; fuscous, nearly straight 
line extending from middle of costa to basal angle of dorsum; at 
apical fourth a transverse, outwardly curved fuscous line extending 
from costa to dorsum ; at end of cell a circle of fuscous scales enclosing 
brown area slightly lighter than rest of wing; entire wing sprinkled 
with occasional fuscous scales; cilia rosy brown. Hindwing whitish 
ochreous with cilia slightly shaded with rose at apex. 

Male genitalia (slide WDD 2095, type) : Uncus bluntly pointed 
apically, slightly recurved; gnathos slightly flattened dorsoventrally ; 
harpe bluntly pointed at apex; anellar lobes symmetrical, broad 
basally, tapering to slightly recurved, acute apex, and with medial 
edges serrate beyond apical third; aedeagus broad basally, narrower 
apically, vesica armed with several large, heavily sclerotized cornuti. 

Female genitalia (slide WDD 3056) : Ductus bursae with patch of 
small spines near inception of ductus seminalis; inception of ductus 
seminalis near corpus bursae. 

Type: In the United States National Museum. 

Type locality: Paraiso, Canal Zone, Panama (S. hovinella); Taboga 
Island, Panama, 200-1000 ft. {S. curiata). 

Distribution : Panama : Paraiso, Canal Zone (June) ; Corozal, Canal 
Zone (June, July); Taboga Island, 200-1000 ft. (Sept.). 

Mejrrick described S. curiata from a series of females and Busck 
described S. hovinella from a series of males; however, examination 
of the types reveals both species identical in color and maculation 
and unquestionably the same species. The failure of past workers 
to accompany descriptions of new species with illustrations has 
resulted undoubtably in many synonyms of this type in the family 
Stenomidae. 

The serrate anellar lobes in the male genitalia and the patch of 
small spines near the inception of the ductus seminalis in the female 
genitalia serve to distinguish L. hovinella from the other species of 
Leihata. 

Lethata buscki, new species 

Figures le,f, 4d 

Alar expanse 19-21 mm. 

Antenna whitish basally, brown beyond. Head whitish shaded 
with rose; second segment of labial palpus brown on outer side, 



NEOTROPICAL MICROLEPIDOPTERA — DUCKWORTH 103 

whitish on inner side, apical segment whitish. Legs whitish, tarsi 
shaded with brown. Thorax brownish shaded with rose. Forewing 
deep ochreous with costa rose; spot at end of cell composed of ring 
of fuscous surrounding white scales; cilia rose basally, whitish beyond. 
Hindwings light ochreous slightly overcast with gray scales; cilia 
light ochreous tinted with rose. 

Male genitalia (slide WDD 3048, type) : Uncus bluntly pointed, 
slightly recurved; gnathos short; harpe of approximately imiform 
width from base to apical third, apex bluntly pointed; anellar lobes 
symmetrical, broad basally, narrowing sharply at middle forming 
narrow, apically acute projections; aedeagus short, apex with large 
hooked process, vesica armed with several heavy cornuti. 

Female genitalia: Unloiown. 

Type: Punta Gorda, British Honduras. USNM 66685. 

Distribution: British Honduras: Punta Gorda (July). Honduras: 
Rio Temas. 

Described from male holotype and one male para type, July 1933 
(G. H. and J. L. Sperry), Punta Gorda, British Honduras; and four 
male paratypes, Rio Temas, Honduras. 

The large hooked process on the apex of the aedeagus and the 
shape of the anellar lobes in the male genitalia readily distinguish 
this species from all others. 

Lethata fusca, new species 

Figures lg,h, Ae 

Alar expanse 25 nmi. 

Antenna brownish. Head gray shaded with light purple, w^hitish 
between eyes; second segment of labial palpus fuscous on outer side, 
whitish on inner side, apical segment whitish. Legs whitish ochreous, 
forelegs and tarsi of mid- and hindlegs heavily shaded with fuscous. 
Thorax fuscous slightly shaded with purple, with median posterior 
tuft of white-tipped fuscous scales. Forewing fuscous with slight 
purplish hue; costa narrowly light rosy ochreous; dark fuscous dot 
in fold at basal third; spot at end of cell white edged with dark 
fuscous; outwardly curved, transverse row of dark fuscous dots at 
apical four-fifths; cilia fuscous. Hindwing whitish ochreous shaded 
with gray; cilia whitish ochreous. 

Male genitalia (slide WDD 3054, type) : Uncus bluntly pointed, 
slightly recurved; gnathos short; harpe narrow, apex pointed; anellar 
lobes symmetrical, roughly triangular in shape with a large spine at 
apex; aedeagus long, narrow, vesica without cornuti. 

Female genitalia: Unknown. 



VOL. 119 



104 PROCEEDINGS OF THE NATIONAL MUSEUM 

Type: Santarem, Brazil. USNM 66686. 

Distribution: brazil: Santarem. 

Described from male holotype, 7.27. (Z.), Santarem, Brazil. 

The specunen from which this species is described bears labels 
indicatmg it had been compared with the type of Stenoma asthenopa 
with which It agi-ees in color and maculation but in a darker shade. 
Examination of the genitalia indicates that the specimen is not L 
asthenopa but rather a new species. The apical spine on the anellar 
lobes and the absence of cornuti in the aedeagus of the male genitalia 
readily separates L. fusca from L. asthenopa and the other species of 
Lethata. 

Lethata glaucopa (Meyrick), new combination 

FiGUEES la, 2a, 4/ 

Stenoma glaucopa Meyrick, 1912, Trans. Ent. Soc. London, vol. 1911, p. 712. 

Alar expanse 32-38 mm. 

Antenna whitish basaUy, darker beyond. Head rosy fulvous 
lighter between eyes; second segment of labial palpus brown, apical 
segment whitish. Legs whitish shaded with brown. Thorax brown 
shaded with rose. Forewing brownish shaded with yellow with costa 
narrowly rosy to brick red; spot at end of cell whitish with dark 
gray center; cilia brown. Hindwing whitish, shaded with brown to 
dark brown; cilia brown. 

Male genitalia (sHde WDD 2684): Uncus pointed, recurved- 
gnathos short, broad at tip; harpe bluntly pointed at apex; anellar 
lobes symmetrical, short, broad, nearly truncate at apex; aedeagus 
large with slight lateral bend, apex with two laterally du-ected processes 
at apex, one extending from each side, the larger one slightly recurved 
vesica without cornuti. ' 

Female genitalia: Illustrated in Clarke (1955, p. 313, figs. 4-4c). 
The illustration is not clear enough to provide a detailed description 
of the female genitalia. 

Type: In the British Museum (Natural History). 

Type locality: San Antonio, Colombia, 5800 ft. 

Distribution: Colombia: San Antonio, 5800 ft. (Nov.). 

This is the largest species in the genus Lethata. It is known only 
from the type, a female in the British Museum, and a male specimen 
m the U.S. National Museum, both from the same locality. The 
two laterally directed apical processes on the aedeagus in the male 
genitalia readily separates L. glaucopa from the other species. 



NEOTROPICAL MICROLEPIDOPTERA — DUCKWORTH 105 

Lethata invigilans (Meyrick), new combination 

Figures 2b,c, 36, 5a 

Stenoma invigilans Meyrick, 1915, Exotic Microlepidoptera, vol. 1, p. 476. 

Alar expanse 24-26 mm. 

Antenna bro\vnish. Head whitish; second segment of labial palpus 
brown on outer side, whitish on inner side; apical segment whitish. 
Legs whitish, heavily shaded with brown on outer surfaces. Thorax 
light brown. Forewing light brown with scattered fuscous scales; 
costa dull rose; three indistinct fuscous transverse lines, one at basal 
thhd, one at middle, and one at apical third; large, conspicuous 
fuscous spot at end of cell; marginal series of fuscous dots around apex 
and termen; cilia whitish gray slightly shaded with rose. Hindwing 
gray; cilia pale grayish. 

Male genitalia (slide WDD 2803): Uncus pointed, recurved; 
gnathos very long, finger-like; harpe narrow, slightly recurved, pointed 
at apex; anellar lobes sj'mmetrical, of near uniform width throughout, 
platelike, only slightly recurved, aedeagus large, broad basally, anterior 
lip of apex somewhat produced, vesica armed with ladder-like series 
of large to small cornuti extending from base to apex. 

Female genitalia (slide WDD 2809) : Ductus bursae with corrugated, 
ribbon-like sclerotized area from ostium to inception of ductus 
seminalis; inception of ductus seminalis near corpus bursae. 

Type: In the British Museum (Natural History). 

Type locality: Maroni River, French Guiana. 

Distribution: French guiana: Maroni River, Caj^enne. Surinam: 
Cottica River, Moengo (May). Venezuela: Sanariapo, Amazonas 
(Aug.). 

For many years this species has been known only from the two 
male specimens on which Meyrick based the original description. 
During the course of the present study, four additional specimens, 
three males and one female, were discovered among the unidentified 
material in the U.S. National Museum collection. These specimens 
not only broadened our knowledge of the distribution of the species 
but also allowed description of the female, which was not previously 
laio wn . 

The large, conspicuous, fuscous spot at the end of the cell on the 
forewing provides the most distinct superficial character for recognition 
of this species; however, one specimen studied had this spot consider- 
ably reduced, suggesting that it is more variable than previously 
thought and should be used with reservations. The ladder-like 
arrangement of the cornuti in the aedeagus of the male genitalia and 
the schlerotized ribbon-like ai-ea extendmg from the ostium to the 



106 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

inception of the ductus seminalis in the female genitalia readily 
identifies this species. 

Lethata leucothea (Busck), new combination 

Figures 2d, 56 

Stenoma leucothea Busck, 1914, Proc. U.S. Nat. Mus., vol. 47, p. 50. 

Alar expanse 20 mm. 

Antenna light bro\vn. Head rosy ochreous; second segment of 
labial palpus brown on outer side, pale rosy ochreous on inner side; 
apical segment pale rosy ochreous. Legs whitish, forelegs heavUy 
shaded with brown, mid- and hindlegs slightly shaded with brown. 
Thorax rosy ochreous, with posteromedian ridge of raised reddish- 
brown scales. Forewing pale ochreous gray with costa narrowly 
brick red; dorsal edge narrowly deep ochreous; spot at end of cell 
consisting of light yellow spot ringed with fuscous scales; outwardly 
curved transverse fuscous line at apical third; less distinct transverse 
fuscous line at middle; an even less distinct transverse line at basal 
third; cilia rosy ochreous. Hindwing light ochreous shaded with 
gray scales; cilia grayish ochreous. 

Male genitalia (slide AB, type): Uncus recurved at tip; gnathos 
short, broad; harpe short, of approximately equal width throughout, 
broadly rounded at apex; anellar lobes symmetrical, bluntly pointed 
at apex; aedeagus large, robust, vesica armed with three large, heavily 
sclerotized cornuti. 

Female genitalia : Unknown. 

Type: In the United States National Museum. 

Type locality: Trinidad Kiver, Panama. 

Distribution: Panama: Trinidad River (March). 

This species is Imown only from the type specimen. It is closest 
to L. hovinella; however, the shape of the harpes and anellar lobes, 
together with the number of cornuti in the aedeagus of the male 
genitalia, serve to separate the two species. 

Lethata maculata, new species 

Figures 2e,f, 3c, 5c 

Alar expanse 24-31 mm. 

Antenna brownish. Head whitish ochreous shaded with rose; 
second segment of labial palpus brown on outer side, whitish on inner 
side, apical segment white on outer side, brown on inner side. Legs 
whitish shaded with brown. Thorax deep yellow ochreous slightly 
shaded with rose, with tuft of white-tipped gray scales posteriorly. 
Forewing deep yellow ochreous with costa narrowly red ochreous; 



NEOTROPICAL MICROLEPIDOPTERA — DUCKWORTH 107 

area of faint rose runs parallel to, and just below, costa; dorsum 
narrowly reddish brown; indistinct fuscous spot on costa at middle, 
from which indistinct, oblique fuscous line runs to anal angle; in- 
distinct, outwardly curved, transverse fuscous line at apical four- 
fifths; spot at end of cell a rather indistinct area of white surrounded 
by fuscous scales; apical margin rich yellow; entire wing sprmkled 
with fuscous scales; cilia rosy ochreous. Hindwing ochreous, anal 
area shaded with gray; cilia rosy ochreous. 

Male genitalia (slide WDD 3061, type): Uncus bluntly pointed, 
slightly recurved; gnathos long, finger-like; harpe short, broadest near 
middle, bluntly pointed at apex; anellar lobes elongate triangular- 
shaped, pointed at apex; aedeagus small, narrow, vesica armed with 
two clusters of small cornuti at apex and single, large cornutus basally. 

Female genitalia (slide WDD 2799, paratype): Ductus bursae with 
corrugated, pouchlike area just before ostium bursae; inception of 
ductus seminalis near ostium. 

Type: Nova Teut6nia, Brazil, 3500 m. USNM 66687. 

Distribution: brazil: Nova Teutonia (Oct.). Surinam: Cottica 
River, Moengo (May). 

Described from male holotype, Oct. 21, 1948 (Fritz Plaumann), 
Nova Teutonia, Brazil, 3500 m.; one male paratype, 1.20 (Parish), 
Tefe, Brazil; and one female paratype, Oct. 21, 1948 (Fritz Plaumann), 
Nova Teutonia, BrazU, 3500 m. 

This species is distinguished readily by the two apical clusters of 
small cornuti and a single, large basal cornutus in the aedeagus of the 
male genitalia and by the ductus bursae of the female genitalia with 
a corrugated, pouchlike area near the ostium bursae. 

Lethata pyrenodes (Meyrick), new combination 

Figures 2g, 5d 
Stenorna pyrenodes Meyrick, 1915, Exotic Microlepidoptera, vol. 1, p. 448. 

Alar expanse 19 mm. 

Antenna brownish. Head, palpus, and thorax deep ochreous shaded 
with rose. Forewing yellow ochreous, fuscous triangular spot at 
middle of costa; spot at end of cell small, fuscous; faint fuscous line 
from costal spot to dorsum; outwardly curved, transverse fuscous 
line from apical third of costa to near tornus; cilia whitish ochreous 
shaded with rose. Hindwings whitish ochreous, apex and termen 
faintly yellowish; cUia whitish ochreous. 

Male genitalia (slide WDD 2817) : Uncus pointed, sharply recurved; 
gnathos short, broad; harpe short, broad, rounded at apex; anellar 



VOL. 116 



108 PROCEEDINGS OF THE NATIONAL MUSEUM 

lobes symmetrical, approximately same width throughout; aedeagus 
large, with large, heavUy sclerotized process posteriorly, vesica with- 
out cornuti. 

Female genitalia: lUustrated in Clarke (1955, p. 350, figs 4-4c) 
The illustration is not clear enough to provide a detailed description 
ol the female genitalia. 

Type: In the British Museum (Natural History). 

T5^pe locaHty: Parang, Argentina. 

Distribution: Argentina: Parana, brazil: Castro, Parana. 

This species is known only from the female type in the British 
Museum and one male specimen in the U.S. National Museum The 
short broad harpe and large, heavily sclerotized, posterior process in 
the aedeagus of the male genitalia readily separates L. pyrenodes from 
ail other species. 

Lethata ruba, new species 

Figures 3a, 5e 
Alar expanse 30 mm. 

Antenna whitish basally, brown beyond. Head whitish shaded with 
rose, face hghter; second segment of labial palpus brown on outer side 
hghter on inner side, apical segment brown. Legs whitish shaded 
with brown. Thorax deep yellow ochreous lightly shaded with rose 
with slight tuft of gray scales posteriorly. Forewing deep yellow 
ochreous with costa narrowly brick red; broad band of pink extends 
parallel to costa from base to apex blending into groundcolor at 
apical third; spot at end of cell fuscous; few fuscous scales scattered 
over entire wing; ciHa whitish tinted with rose. Hindwing whitish 
ochreous lightly shaded with gray; ciha whitish. 

Male genitaHa (slide WDD 2S00, type): Uncus very short, broad 
sharply recurved; gnathos short, V-shaped at tip; harpe short, rounded 
at apex; anellar lobes asymmetrical; one narrow, pointed at apex; one 
broad, with three large spines apically. Aedeagus with apex on one 
side produced into long bladelike process, vesica without cornuti. 

Female genitaha: Unknown. 

Type: Nova Teutonia, Brazil, 3500 m. USNM 66688. 

Distribution: brazil: Nova Teutonia, 3500 m. (Aug.). 

Described from male holotype, Aug. 6, 1948 (Fritz Plaumann). 
Nova Teutonia, Braz^, 3500 m. 

The shape of the asymmetrical anellar lobes and the bladelike 
process on the apex of the aedeagus distinguish this species from all 
others in the genus Lethata. 



NEOTROPICAL MICROLEPIDOPTERA — DUCKWORTH 109 

Lethata aatyropa (Meyrick), new combination 

Figures 2h-j, 5/ 
Stenoma satyropa Meyrick, 1915, Exotic Microlepidoptera, vol. 1, p. 478. 

Alar expanse 24-25 mm. 

Antenna brownish. Head rosy whitish, crown suffused with deep 
ochre; second segment of palpus brown on outer side, white on inner 
side, apical segment whitish. Legs whitish shaded with brown. 
Thorax deep ochreous, becoming darker posteriorly. Forewing deep 
ochreous with costa dull red from base to apical third; dorsum nar- 
rowly brown red throughout; an indistinct dark line at basal third 
from dorsum to middle, a large, round fuscous spot at end of cell 
containing transverse purplish spot; faint, curved, transverse line 
from costa to dorsiun at apical four-fifths ; entire wing sprinkled with 
fuscous scales; cilia ochreous, suffused with purplish on outer half, 
Hindwings whitish ochreous slightly overcast with gray scales, apical 
and terminal edge yellow; cilia white, tinted with rose. 

Male genitalia (slide WDD 2651): Uncus pointed apically, slightly 
recurved; gnathos short, blunt at apex; harpe broadest at midpoint, 
bluntly pointed at apex; anellar lobes symmetrical, apex of each 
developed into strongly recurved, hooklike spine; aedeagus tapering 
from middle to extremely sharp-pointed apex, single long spine projects 
posteriorly from near middle, vesica armed with numerous spiculate 
cornuti. 

Female genitalia: Unknown. 

Type : In the British Museum (Natural History) . 

Type locality: Godebert, Maroni River, French Guiana. 

Distribution: French guiana: Godebert, Maroni River. 

This species is known only from the male type in the British Museum 
and a male specimen in the U.S. National Museum, both from the 
same locahty. The shape of the aedeagus and the large posterior 
process from its middle, plus the shape of the anellar lobes, provide 
ample distinguishing characters in the male genitalia. 



Literature Cited 

Clarke, J. F. Gates 

1955. Catalogue of the type specimens of Microlepidoptera in the British 
Museum (Natural History) described by Edward Meyrick, vol. 2. 



no 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figure 1. — Lateral view of abdominal segments 1-3: a, Lethata glaucopa (Meyrick). 
Ventral view of male genitalia, aedeagus removed: b, L. trochalosticta (Walsingham); 
f, L. buscki, new species; g, L. fusca, new species; j, L. bovinella (Busck); k, L. aromatica 
(Meyrick). Aedeagus: c, L. trochalosticta (Walsingham); e, L. buscki, new species; 
h, L. fusca, new species; i, L. bovinella (Busck); 1, L. aromatica (Meyrick). Wing vena- 
tion: d, L. trochalosticta (Walsingham). 



NEOTROPICAL MICROLEPIDOPTERA — DUCKWORTH 



111 








Figure 2. — Ventral view of male genitalia. Aedeagus in situ : a, Lethata glaucopa (Meyrick) ; 
d, L. leucothea (Busck); g, L. pyrenodes (Meyrick); h, L. saiyropa (Meyrick). Aedeagus 
removed: b, L. invigilans (Meyrick); e, L. maculata, new species. Aedeagus: c, L. 
invigilans (Meyrick) ; f , L. maculata, new species. Lateral view of aedeagus : i, L. satyropa 
(Meyrick). Enlarged view of bifurcate seta: j, L. satyropa (Meyrick). 



112 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 




Figure 3.— Ventral view of male and female genitalia: a, Lethata ruba, new species, aedeagus 
in situ; b, L. invigilans (Meyrick); c, L. maculate, new species; d, L. bovinella (Busck). 



U.S. SOVERNMENT PRINTING OFPICEil964 



NEOTROPICAL MICROLEPIDOPTERA DUCKWORTH 113 




"iGURE 4. — Left wing: a, Lethala truchalosticta (Walsingham); b, L. aroinatica (Mcyrick); 
c, L. bovinella (Busck); d, L. buscki, new species; e, L. fusca, new species; f, L. glaucopa 
(Mcyrick). 



rL'T M?> 04 



114 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 




Figure 5. — Left wing: a, Lethata invigilans (Meyrick); b, L. leucuthea (Busck); c, L. mac- 
ulata, new species; d, L. pyrenodes (Meyrick); e, L. ruba, new species; f, L. salyropa 
(Meyrick). 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1964 Number 3498 



ONE NEW SPECIES AND TWO REDESCRIPTIONS OF 
CATFISHES OF THE SOUTH AMERICAN CALLICHTHYID 
GENUS CORYDORAS 



By Stanley H. Weitzman 



Recent examination of three specimens of a new catfish from 
Brazil, herein described as Corydoras semiaquilus, has instigated re- 
examination and redescription of its apparent relatives, C. treitlii 
Steindachner (1906) and C.fowleri Bohlke (1950). C.fowleri is known 
only from the holotype, which was described without illustration. In 
addition, the original description of C. fowleri, like that of C. cochui 
Myers and Weitzman (1954) (see Weitzman, 1956), was published in 
an aquarium magazine not available in many university libraries, 
C. treitlii has never been illustrated and its original description, al- 
though excellent for its time, is now inadequate in the light of many 
subsequent new species. Discovery of apparent close relatives of C. 
treitlii has made reexamination and description of the type specimens 
imperative. 

I am indebted to Professor George S. Myers for the loan of speci- 
mens and the use of facilities in the Division of Systematic Biology 
at Stanford University ; to Dr. Paul Kahsbauer of the Naturhistorisches 
Museum, Wien, for the loan of type specimens of Corydoras described 
by Franz Steindachner; to Mr. W. I. Follett, Curator of Fishes of the 
California Academy of Sciences, for the loan of specimens; to Mr. 

115 



116 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Harald Schultz, Sao Paulo, Brazil, for locality data and for collecting 
the specimens of C. semiaquilus; and to Dr. Herbert Axelrod of Jersey 
City, New Jersey, for two specimens of C. semiaquilus. I also wish 
to thank Dr. W. Klausewitz and Mr. Fritz Rossel of the Sencken- 
bergische Naturforschende Gesellschaft, Natm"-Museum und For- 
schungs-institut, for the loan of a specimen of C. semiaquilus. 
The following abbreviations are used: 

NMW — Naturhistorisches Museum, Wien 
USNM — United States National Museum 
SU — Division of Systematic Biology, Department of Biological Sciences, 

Stanford University 
SM — Senckenberg Museum. 

Corydoros treitlii Steindachner 

Figures 1 and 2 

Corydoras treitlii Steindachner, 1906, p. 478 (original description; type locality: 
mouth of a small stream emptying into the Rio Parnahyba at Victoria, State 
of Maranhao, Brazil). — Eigenmann, 1910, p. 403 (listed). — Miranda Ribeiro, 
1911, p. 167 (description copied from Steindachner, 1906). — Regan, 1912, p. 
210 (description copied from Steindachner, 1906). — Ellis, 1913, p. 407 
(listed). — Gosline, 1940, p. 15 (aquarium specimen, no description) ; 1945, p. 
74 (listed). — Stigchel, 1946, p. 129 (description of specimen from original 
collection). — Bohlke, 1950, p. 27 (discussion of relationships with Corydoras 
/oa'?m).— Fowler, 1954, p. 67 (listed). 

Lectotype. — NMW 61103, standard length 42.6 mm., collected 
during 1903 by Franz Steindachner at mouth of brook emptying into 
Rio Paranhyba [Parnaiba River] at Victoria [Alto Parnaiba], State 
of Maranhao, Brazil. 

Additional specimens. — NMW 47798, paralectotype, standard 
length 42.4 mm., same data as lectotype; USNM 176912, standard 
length 52.5 mm., Sao Paulo, Brazil, Herbert Axelrod, 1958; SU 35054, 
standard length 47.0 mm., sent to the Division of Systematic Biology, 
Stanford University, by Mr. Fred H. Stoye in March 1937. Mr. 
Stoye stated that, according to Mr. N. Greim, this is an aquarium 
specimen from the Amazon. In my opinion this locality data is 
uncertain. This is the specimen utilized by Gosline (1940). 

Diagnosis. — Corydoras treitlii may be distinguished from other 
species of Corydoras by the following combination of characters: 
Snout long, about 30 to 32% of body length mthout head. Least 
caudal peduncle depth about 57 to 62% of snout length. Imbricated 
thoracic and abdominal plates absent; fine bony pricldes present 
in these regions. Dorsal fin spine about equal in length to pectoral 
fin spine. Predorsal length about 79 to 89% of distance between 
dorsal fin origin and caudal fin base. Caudal fin without bars. 



CATFISHES OF THE GENUS CORYDORAS — WEITZMAN 



117 



Description. — (For actual measurements see table 1.) In the 
description below, the proportions are given first, percentages follow 
in parentheses, both of which derive from standard length unless 
otherwise designated. Data for the lectotype, NMW 61103, is 
given fii-st, data for NMW 4779S follows in brackets. Data for USNM 
179612 and for SU 35054 are designated by respective abbreviations. 




Figure 1. — Corydoras ireillii Steindachner, lectotype, NMW 61103. Standard length 

42.6 mm. 



Body fairly elongate, compressed posteriorly. Greatest body depth 
2.9 (34.8%) [2.8 (35.6%)], USNM 2.8 (35.8%), SU 2.9 (34.4%). 
Least depth of caudal peduncle 7.5 (13.4%) [7.2 (13.9%)], USNM 7.7 
(15.0%), SU 7.6 (13.1%). Distance between snout tip and dorsal fin 
origin 1.8 (56.4%) [2.0 (49.1%)], USNM 2.0 (51.0%), SU 1.9 (52.3%). 
Distance between snout tip and anus 1.9 (52.8%) [1.8 (55.0%)], 
USNM 1.9 (52.0%), SU 2.0 (49.8%). Anal fin origin to snout tip 
1.3 (80.0%) [1.2 (80.8%)], USNM 1.3 (79.8%), SU 1.2 (80.6%). 
Lateral scutes 24/21 in all four specimens. Abdomen and thorax 
with small prickles in all specimens, no heavy imbricated plates. 
In SU 35054, a poorlj'- preserved specimen, many of these pricldes have 
been broken off in exposed areas but their bases remain. Azygous 
middorsal scutes 6 [4], USNM 4, SU 5, before adipose fin. One 
ar.ygous scute before dorsal fin in all specimens. Pectoral fin base 
incompletely surrounded by coracoid in all specimens. Distance 
between coracoids variable (probably ■wider in females than in males) 
10.9 (9.2%) [9.2 (10.9%)], USNM 11.3 (8.8%), SU 13.4 (7.5%). 
Head length 3.0 (33.3%) [2.9 (33.8%)], USNM 3.0 (33.3%), SU 
(34.2%). Greatest head width 1.4 (71.8%) [1.5 (68.7%)], USNM 1.5 
(67.4%), SU 1.7 (59.5%) in head length. Least width of bony inter- 



118 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

orbital 3.2 (31.0%) [4.0 (25.2%)], USNM 3.1 (32.0%), SU 3.5 
(28.4%) in head length. Snout acute in dorsal view. Snout tip 
rounded in lectotype, much more acute in SU 35054 (a poorly pre- 
served, dehydrated specimen). Snout 1.4 (70.4%) [1.5 (66.4%)], 
USNM 1.6 (64.0%), SU 1.5 (64.9%) in head length. Dorsal profile 
of snout slightly concave in all specimens. When directed posteriorly, 
both upper rictal (actually maxillary) and lower rictal barbels reach 
a point on a vertical about half an orbital diameter behind posterior 
edge of the orbit. Greatest diameter of bony orbit 3.8 (26.1%) 
[4.6 (21.7%)], USNM 4.1 (24.6%), SU 3.9 (25.8%) in head length. 
Greatest width of suborbital 2.3 (43.3%) [1.5 (67.7%)], USNM 1.4 
(69.8%), SU 2.4 (41.1%) in orbit. 

Dorsal fin I, 7, last fin ray split to its base in all specimens. Spine 
of dorsal fin when depressed reaching to, or slightly beyond, posterior 
termination of dorsal fin base, distant from origin of adipose fin. 
Adipose fin spine in orbit 1.0 (97.4%) [1.0 (100.3%)], USNM 
1.3 (79.0%), SU 1.2 (84.7%). Anal fin ii,5, last ray split to its base 
in all specimens. USNM 179612 could be interpreted as ii,6, last 
fin ray not split to its base. The last two ray elements of the anal 
fin in this specimen are well separated and probably each ray base 
belongs to its own separate pterygiophore series. Pectoral fin 
1,10, [1,11], USNM 1,10, and SU 1,10. Pelvic rays i,5, in all speci- 
mens. Principal caudal rays 7/7 in all specimens. Pectoral fin 
spine (see fig. 2) with 16 [18], USNM 21, SU 16, stout spinules along 
its posterior border. 




Figure 2. — Corydoras treitlii Steindachner. Pectoral fin spine of lectotype, ventral 

view, left spine. 

Color. — The lectotype has the following color pattern in alcohol 
(see also fig. 1): Basic body color yellowish gray. Head with brown 
marldngs as shown in figure 1 ; no punctate or vermiculate markings 
over snout or head. Sides with purplish dark brown markings on 
upper body scutes. All specimens with all fins hyaline, completely 
lacking bars, bands, or blotches. All specimens with color pattern 
very similar to lectotype except that SU 35054 greatly faded. Color 
in life not known. 



CATFISHES OF THE GENUS CORYDORAS — WEITZMAN 119 

Corydoras fotcleri Bohlke 

Figures 3 and 4 

Corydoras foiuleri Bohlke, 1950, p. 2G (original description; type locality: Cano 
del Chancho, near Pevas, Peru). 

Holotype: SU 16115, a female 66.0 mm. in standard length, 
collected December 13, 1941, by Mr. William G. Scherer at Cafio 
del Chancho, near Pevas [Pebas], Peru. 

Diagnosis. — Corydoras fowleri may be distinguished from other 
known species of Corydoras by the following combination of characters: 
Snout relatively long, about 25% of body length without head. 
Least caudal peduncle depth about 55% of snout length. Imbricated 
thoracic and abdominal plates present. Dorsal fin spine weaker and 
shorter than pectoral fin spine. Predorsal length about 67% of 
postdorsal length. 




Figure 3.— Corydoras fowleri Bohlke, holotype, SU 16115. Standard length 66.0 mm. 

Description. — (See table 1 for measurements.) For explanation 
of descriptive pattern below, see p. 117. 

Body elongate, especially slender posterior to dorsal and pelvic fins; 
greatest body depth 3.3 (30.8%). Least depth of caudal peduncle 10.0 
(10.0%). Dorsal fin origin much nearer to snout tip than caudal 
fin base. Distance between snout tip and dorsal fin origin 
2.3 (43.4%). Distance between snout tip and anus 2.0 (49.5%). 
Anal fin origin to snout tip 1.2 (80.0%). Lateral scutes 24/21. 
Abdomen and thorax entirely covered with small- to moderate-sized 
imbricated plates. Azygous middorsal scutes 5 anterior to adipose 
fin, 1 anterior to dorsal fin. Pectoral fin base incompletely sur- 
rounded by coracoid, interval or hiatus filled with moderate-sized 
plates continuous with those of abdomen and thorax. Distance 
between coracoids 7.8 (12.9%). Head length 3.8 (26.6%); gi-eatest 



120 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

head width 1.3 (75.5%) in its length. Least width of bony inter- 
orbital 3.1 (32.0%) in head length. Snout acute in dorsal view but 
snout tip rather rounded. Snout length 1.4 (69.2%) in head length. 
Dorsal profile of snout slightly concave. Wlien directed posteriorly, 
both rictal barbels of both sides reach a point on a vertical line from 
posterior eye margin. Greatest diameter of orbit 4.4 (22.8%) in 
head length. Greatest width of suborbital 2.9 (35.0%) in orbit. 
Dorsal fin 1,7, last fin ray split to its base. Dorsal fin damaged, 
its spine and anteriormost rays partially lost as shown in figure 3. 
Dorsal fin spine rather slender, considerably more slender than the 
pectoral fin spines. Adipose fin spine 0.95 (104.8%) in orbit. Anal 
fin ii,5, last ray split to its base. Bohlke (1950) reported the anal 
fin count as "1,5, the last ray widely split to its base." I found 
that in taking counts and measurements I had counted this fin as 
i,5; however, during careful examination of the fin while preparing 
figure 3, I found that what appeared to be a single nonpungent spine 
was actually two very closely adpressed nonpungent spines. Pectoral 
elements 1,10. Pelvic fin rays i,5 on both sides. Caudal fin, although 
both lobes broken, with principal rays 7/7. Pectoral fin spine (fig. 4) 
has 30 spinules along its posterior border. 




Figure 4. — Corydoras foivleri Bohlke. Pectoral fin spine of holotype, ventral view, left 

spine. 

Color. — Specimen appears somewhat faded. Overall body color 
very pale brown. Ventral region below gills white. Just posterior to 
this region, at origin of belly scutes, belly region becomes very pale 
brown. Pattern of dark brown markings shown in figure 3. This 
pattern described by Bohlke (1950) as follows: 

. . . dark blotch along base of dorsal fin, abruptly shifted ventrally at a 
vertical with the fifth articulated dorsal ray. This posterior continuation 
of the blotch covers the lower two-thirds of the upper row of lateral scutes, 
and continues back to below the adipose dorsal. Several small dark spots at 
the upper end of the gill openings, somewhat resembling a broken-up humeral 
spot. Top of head, snout, and cheeks covered by wav}' longitudinal dark 
lines. 

Little can be added to this description. 

Discussion. — As noted by Bohlke (1950), Corydoras fowleri appears 
to be related to C. treitlii; however, as he remarked, it differs from 0. 
treitlii in possessing imbricated thoracic and abdominal plates and a 
longer body in relation to head length. In addition to these charac- 



CATFISHES OF THE GENUS CORYDORAS — WEITZMAN 121 

ters, it has a much weaker (and probably shorter) dorsal fin spine than 
pectoral spine. The pectoral and dorsal fin spines of C. treitlii are 
about equal in diameter and length, Corydoras fouieri appears closely 
related to C. semiaquilus but differs in the characteristics noted 
under the latter species. 

The swollen appearance of the holotj^pe of Corydoras fouieri indicates 
that the specimen is probably a gravid female. This probabihty was 
verified by probing between the lower third and fourth lateral scutes 
of the right side in the upper regions of the coelomic cavity, where 
a few mature eggs were recovered. Since the holotype and only known 
specimen of C. jowleri is a female, apparently swollen with ripe eggs, 
and since males of the genus Corydoras are usually slenderer than 
their female counterparts, it is quite Hkely that the species C. jowleri 
has an average body depth much less than that indicated here. Thus, 
C. fouieri probably differs more from C. semiaquilus and C. treitlii 
with regard to body depth than the data avilable at present would 
indicate. 

In summary, Corydoras foioleri is related most closely to C. semi- 
aquilus but differs from that species in its shorter snout, longer caudal 
peduncle length, lesser body depth, proportionately smaller eye, 
and shorter head length. 

Corydoras semiaquilus, new species 

Figures 5 and 6 

Holotype. — ^SU 55939, standard length 60.9 mm., collected during 
December of 19G0 by Harald Schultz from Igarape Preto, according 
to Mr, Schultz, "a small jungle-rivulet at the headwaters of the black- 
water creeks, which empty in the upper Solimoes, State of Amazonas, 
Brazil. These small and narrow creeks have crj'-stal clear water, 
sandy bottom, covered with pebbles, leaves and rotten leaves." 

Additional specimens. — USNM 196170, standard length 59.1 mm.; 
SM 5349 damaged but standard length about 54 mm. Both with 
the same data as the holotype. 

Diagnosis. — Corydoras semiaquilus may be distinguished from 
other species of Corydoras by the following combination of characters: 
It has a long snout (snout length about 32 to 33% of body length 
without head). Least caudal peduncle depth about 49 to 51% of 
snout length. Most species of Corydoras, except for the long snouted 
species, have the least depth of the caudal peduncle and the snout 
length about equ-al. Imbricated thoracic and abdominal plates pres- 
ent. Dorsal fin spine considerably weaker and shorter than pectoral 
fin spine. Predorsal length about 85 to 86% of distance between 
dorsal fin origin and caudal fin base. Caudal fin heavily barred. 



122 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Description. — ^(See table 1 for measurements.) For explanation 
of descriptive pattern below, see p. 117. SU 55939 is given first, 
USNM 196170 follows in brackets. Measurements of SM 5349 are 
not given because the head is badly damaged, making accurate 
measurements impossible. 




Figure 5. — Corydoras semiaquilus, new species, holotype, SU 55939. Standard length 

60.9 mm. 



Body relatively elongate, greatest body depth 3.0 (33.5%) [3.0 
(33.5%)]. Least depth of caudal peduncle 8.9 (11.2%) [8.6 (11.7%)]. 
Dorsal fin origin nearer snout tij) than caudal fin base. Distance 
between snout tip and dorsal fin origin 2.0 (50.7%) [2.1 (48.2%)]. 
Distance between snout tip and anus 1.9 (52.8%) [1.9 (52.5%)]. 
Anal fin origin to snout tip 1.2 (81.5%) [1.2 (80.4%o)]. Lateral 
scutes 24/21 [26/23]. Abdomen entirely covered with small to moder- 
ate-sized imbricated bony plates. Azygous middorsal scutes 7 [6] 
anterior to adipose fin and 1 [1] anterior to dorsal fin. Pectoral fin 
base incompletely surrounded by coracoid. Distance between 
coracoids 8.5 (11.8%) [9.0 (11.2%o)]- Head length 3.0 (32.9%) 
[3.0 (33.2%)]; greatest head width 1.5 (65.5%) [1.5 (64.8%)] in its 
length. Least width of bony interorbital 4.1 (24.6%) [3.9 (25.5%)] in 
head length. Snout acute in dorsal view but snout tip rather broadly 
rounded. Snout 1.4 (69.5%) [1.4 (69.5%o)] in head length. Dorsal 
profile of snout concave in both specimens. When directed posteri- 
orly, both rictal barbels reach a point on a vertical line passed down 
from posterior eye margin. Greatest diameter of orbit 4.4 (22.5%) 
[4.4 (22.9%o)] in head length. Greatest width of suborbital 2.8 (35.6%) 
[1.8 (55.5%)] in orbit. 



CATFISHES OF THE GENUS CORYDORAS — ^WEITZMAN 123 

Dorsal fin 1,7, in all specimens, last fin ray split to its base. Neither 
first spine nor first soft ray of dorsal fin reaches base of adipose spine 
when dorsal fin depressed. Dorsal fin spine much slenderer and 
shorter than pectoral fin spine. Adipose spine 1.5 (66.7%) [1.3 (77.8%)] 
in orbit. Anal fin ii,5, in all specimens, last ray split to base in SU 
55939 and SM 5349 and not split to its base in USNM 196170. Pec- 
toral fin 1,11, in SU 55939 and USNM 196170, 1,10, in SM 5349. 
Pelvic fin rays i,5, in all specimens. Caudal fin with principal rays 
7/7 in all specimens. Pectoral fin spine (fig. 6) has 26 [24] spinules 
along its posterior border. There are 22 spinules in SM 5349. 




Figure 6. — Corydoras semiaqiiilus, new species. Pectoral fin spine of holotype, ventral 

view, left spine. 

Color. — The holotype has the follo\ving color in alcohol (see also 
fig. 5): Ground color white to pale brown. Head with punctate 
to vermiculate dark brown or black markings, more punctate over snout 
and between eyes. Sides of head with black to gray markings. Belly, 
thorax, and lower one-third of body white. Upper two-thirds of body 
dark except for narrow streak of pale brown below dorsal fin. This 
pale area extends posteriorly to caudal fin above dorsal margin of 
body. Dorsal fin with two rather irregular black bars; caudal fin 
with thi-ee kregular black vertical bars. Anal fin with two black 
bars. Pelvic fins colorless except for some scattered melanophores 
over central portions of rays. Basal two-thirds of pectoral fin rays 
dark and entire pectoral fin spine dark. Color of paratypes very 
similar to that of holotype. 

Color in life (from a Kodachrome) as follows: Dark areas seen in 
figure 5 black with slight suggestion of blue and purple. Greenish 
iridescence over lateral part of coracoid bone and golden iridescence 
over opercular bone. Pale areas of body very pale brown, somewhat 
pinkish in thoracic and abdominal regions while pale strip along dorsal 
edge of body somewhat darker brown. Iris around pupU golden, 
shading to black at edge of eye. 

The name "semiaquilus" is from Latin (semi=haK and aquilus = 
dark colored) referring to the dark color on the upper body scutes. 

Discussiox. — The relationships of Corydoras semiaquilus seem to 
be clearly the long-snouted Corydoras such as treitlii, acutus, cervinus, 
fowleri, pastazensis, and septentrionalis. So far as known, it differs 
from all of these but C. fowleri m the possession of a thick laj^er of 
imbricated dermal plates on the abdomen and thorax. The species 



124 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

appears closely related to C. treitlii because of its very long snout, its 
bony interorbital width contained twice or more in the snout length, 
its relatively slender caudal peduncle, and its color pattern. The obvi- 
ous differences between C. treitlii and C. semiaquilus are the imbricated 
bony plates on the belly and thorax of the latter (the former has 
prickles only), a much longer pectoral spine in C. semiaquilus (see 
table 1 for comparison), a banded caudal fin in C. semiaquilus (caudal 
bands are absent in C. treitlii), and fauiy consistent differences in caudal 
peduncle depth between the two species. The profUe of the snout 
and dorsal part of the head is consistently different in C. treitlii and 
C. semiaquilus. Corydoras treitlii has the caudal peduncle depth 57.0 
to 62.1 % of the snout length while in C. semiaquilus it is 48.9 to 50.8%. 

Corydoras cervinus differs from C. semiaquilus in lacldng caudal fin 
bars, in having larger spinules on the pectoral fin spine, and apparently 
in lacking belly scutes. Rossel (1962) records the caudal peduncle 
depth for C. cervinus as 7 times in the standard length. In C semi- 
aquilus it is 8.6 to 8.9 times in the standard length. 

Of the other known species with an interorbital contained twice or 
more in the snout, Corydoras fowleri, 2.2 (42.3%) in snout length, is 
very much like C. semiaquilus in the possession of imbricated thoracic 
and abdominal scutes and a somewhat similar color pattern. How- 
ever, these two species differ widely with regard to several proportions. 
First, C fowleri has a much shorter snout (about 24% of body length 
without head). Second, the head length of C. fowleri is 26.6% and 
that of C. semiaquilus is 32.9 and 33.2% of the standard length. Third, 
the eye of C. fowleri is proportionately smaller, being 6.1%, while that 
of C. semiaquilus is 6.6 to 7.4% of the standard length. Fourth and 
finally, the dorsal fin is placed farther anteriorly in C. fowleri, its pre- 
dorsal length being 66.5% while that of C. semiaquilus is 85.2 to 85.5% 
of the distance between the origin of the dorsal fin and the caudal fin 
base. The color patterns of C. fowleri and C. semiaquilus actually 
may be quite similar; however, that of C. fowleri appears faded and, 
therefore, accurate comparisons cannot be made. 

Another species, Corydoras pastazensis Weitzman (1963), is very 
closely related to C. treitlii and on the basis of specimens at hand, may 
be distinguished from C. semiaquilus by the following characters: 
C. semiaquilus has a body depth of 33.5% of the standard length while 
that of C. pastazensis is 36.4 to 37.1%. Body depth, being measured 
from the ventral border of the posterior coracoid process to the dorsal 
fin origin, is a stable measurement in Corydoras, little affected by the 
state of nutrition. Caudal peduncle depth of C. semiaquilus is 48.9 
to 50.8% in the snout length while it is 65.2 to 66.4% in C. pastazensis. 
Corydoras pastazensis lacks the dark color pattern in the upper body 
scutes present in C. semiaquilus and lacks the marbled markings on 
the snout. 



CATFISHES OF THE GENUS CORYDORAS — WEITZMAN 125 

Table 1. — Measurements of three Corydoras species 







Corydoras treitlii 




Corydoras 
semiaquilus 


Corydoras 
fowleri 


Measurement 


Lecto- 
type 

NMW 
61103 


NMW 
47798 


USNM 
179612 


SU 
35054 


Holo- USNM 
type 196170 
SU 
55939 


Holotype 

SU 

16115 


Standard length 


42.6 


42. 4 


52. 5 


44. 2 


60. 9 59. 1 


66. 


Head length 


14.2 


14.3 


17.5 


15. 1 


20. 19. 6 


17.5 


Snout length 


10.0 


9.5 


11.2 


9.8 


13. 9 13. 6 


12. 1 


Least width of bony inter- 














orbital 


4. 4 


3.6 


5.6 


4.3 


4. 9 5. 


5.6 


Greatest diameter of bony 














orbit 


3.7 


3. 1 


4.3 


3.9 


4. 5 4. 5 


4.0 


Greatest width of suborbital 


1.6 


2. 1 


3.0 


1.6 


1. 6 2. 5 


1.4 


Length of fontanel 


5.5 


5.4 


6.9 


6.5 


6. 2 6. 5 


4.5 


Length of predorsal scale 


2.5 


2.5 


3.4 


3.0 


3. 8 3. 3 


3.7 


Greatest width of head 


10.2 


9.9 


11.8 


9.0 


13. 1 12. 7 


13.2 


Snout tip to dorsal fin origin 


24. 


20.8 


26. 8 


23. 1 


30. 8 28. 5 


28.6 


Snout tip to anal fin origin 


34. 


34.2 


41. 9 


35.6 


49. 7 47. 4 


53.0 


Snout tip to anterior edge 














of anus 


22. 5 


23.3 


27.3 


22.0 


32. 2 31. 


32. 7 


Greatest body depth 


14.8 


15. 1 


18.8 


15.2 


20. 4 19. 8 


20.3 


Least depth of caudal pe- 














duncle 


5.7 


5.9 


6.8 


5. 8 


6. 8 6. 9 


6. 6 


Distance between coracoids 


3.9 


4.6 


4.7 


3.3 


7. 2 6. 6 


8.5 


Length of dorsal spine 


9. 1 


8.6 


10.2 


10.0 


10. 8 10. 6 


? 


Length of pectoral spine 


8.9 


8.5 


11.6 


10. 


14. 2 13. 9 


16.7 


Length of adipose spine 


3.6 


3.2 


3.4 


3.3 


3. 3. 5 


4.2 


Origin of dorsal fin to caudal 














fin base 


27. 1 


26.4 


32. 1 


26.7 


36. 2 34. 5 


43.0 


Post head length 


30.9 


29.9 


37.4 


31.3 


42. 7 41. 5 


49.8 


Caudal peduncle length 


6.3 


6.0 


7.3 


5.9 


8. 5 8. 3 


9.8 



126 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

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1950. A new catfish of the genus Corydoras from the Peruvian Amazon. 
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1910. Catalogue of the fresh-water fishes of tropical and south temperate 

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Ellis, Marion Durbin 

1913. The plated Nematognaths. Ann. Carnegie Mus., vol. 8, nos. 3-4, 
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1954. Os peixes de dgua doce do Brasil (4.» entrega). Arq. Zool. Est. 
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1940. A revision of the neotropical catfishes of the family Callichthj'idae. 
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1911. Fauna Brasiliense: Peixes, IV(A): Eleutherobranchios Aspirophoros. 

Arch. Mus. Nac. Rio de Janeiro, vol. 16, 511 pp., pis. 22-54. 
Myers, George S., and Weitzman, Stanley H. 

1954. Another new Corydoras from Brazil. Aquarium Journ., vol. 25, no. 

4, pp. 93-94. 
Regan, Charles Tate 

1912. A revision of the South American genus Corydoras, with a list of the 

specimens in the British Museum (Natural History). Ann. 
Mag. Nat. Hist., ser. 8, no. 10, pp. 209-220. 
Rossel, Fritz 

1962. Corydoras cervinus, ein neuer Panzerwels aus Brasilien (Pisces, Tele- 

ostei, Calichthyidae [sic]). Senck. Biol., vol. 43, no. 1, jop. 31-33. 
Steindachner, Franz 

1906. Ueber zwei neue Corydoras-Arten aus dem Parnahyba- und Parahim- 
flusse im Staate Piauhy. Anz. Akad. Wiss Wien, vol. 43, no. 27, 
pp. 477-482. 

StIGCHEL, J. W. B. VAN der 

1946. South American Nematognathi, 204 pp., 3 tbls. 
Weitzman, Stanley H. 

1956. A description, supplementary notes and a figure of Corydoras cochui 
Myers and Weitzman, a Brazilian catfish. Stanford Ichth. Bull., 
vol. 7, no. 2, pp. 14-18. 

1963. A new catfish, Corydoras pasiazensis (Callichthyidae), from Ecuador. 
Proc. Biol. Soc. Washington, vol. 76, pp. 59-63. 



U.S. GOVERNMENT PftlNTJNG OFFICE:IS64 



Proceedings of 
the United States 
National Museum 




SMITHSONIAN INSTITUTION . WASHINGTON, D.C. 



Volume 116 



1964 



Number 3499 



OSTEOLOGY AND RELATIONSHIPS OF 

SOUTH AMERICAN CHARACID FISHES OF 

SUBFAMILIES LEBIASININAE AND ERYTHRININAE 

WITH SPECIAL REFERENCE TO SUBTRIBE NANNOSTOMINA ^ 



By Stanley H. Weitzman 



Introduction 



The present work is the result of an attempt to determine the 
relationships of two characid genera, Nannostomus and PoecUobrycon, 
to other members of the cypriniform family Characidae. The two 
genera include about eight known species of small characids found in 
forest brooks of South America. These species are here assigned to 
the subtribe Nannostomina. Although the first known species of 
this group was described in 1872, adequate investigation of morpho- 
logical evidence for their relationships has never been undertaken. 
Previous investigations have been superficial in scope and often 
inaccurate in fact primarily because of the small size of these fishes. 
The total cranial length is usually 5-10 mm, and the largest recorded 
specimen is 44.5 mm. in standard length. In some cases, despite the 
lack of adequate morphological data, certain ichthyologists have 



' This paper is the second of three parts based on a dissertation submitted to 
the Department of Biological Sciences, Stanford University, California, in partial 
fulfillment for the degree of Doctor of Philosophy. See Weitzman (1962) for 
the first part. 

127 



128 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

shown rather good intuitition in their interpretation of the relation- 
ships of these fishes; however, others have not been so fortunate in 
their interpretation and these fishes have been shifted about in the 
classification of characids until it is very difficult to obtain from the 
literature a concept of their affinities. 

Investigation of Nannostomus, Poecilobrycon, and their relatives 
led to comparison of two widely divergent characid subfamilies, the 
Characinae, treated by Weitzman (1962), and the Lebiasininae, 
treated here. In addition the Erythrininae, another divergent 
subfamily of the Characidae, are compared with the Lebiasininae 
because these two groups of characids often have been thought to 
belong to a single group. Although the work began as an effort to 
determine the relationships of Nannostomus and Poecilobrycon, its 
most important result concerns the relationships of the subfamilies 
Lebiasininae and Erythrininae. 

The morphological data obtained in the present study has resulted 
in the following classification, the names given below being used 
throughout this paper (see pages 148 to 152 for a full treatment of the 
classification) : 

Subfamily Lebiasininae 

Tribe Lebiasinini 
Tribe Pyrrhulinini 

Subtribe Pyrrhulinina 

Subtribe Nannostomina 
Subfamily Erythrininae 

For the loan of specimens, I am indebted to Dr. George S. Myers 
of Stanford University, Mr. W. I. Follett of the California Academy 
of Sciences, Dr. James Bohlke of the Academy of Natural Sciences 
of Philadelphia, and Mr. Loren P. Woods of the Chicago Natiu-al 
History Museum. I am indebted especially to Dr. George S. Myers 
for critically reading much of the manuscript and offering aid and 
advice during most of its preparation. In addition, the following 
persons have read the manuscript at various stages, all providing very 
useful help: Mrs. LiUian Dempster of the California Academy of 
Sciences, Drs. Myra Keen and Warren Freihofer of Stanford Univer- 
sity, and Drs. Leonard P. Schultz, Kobert H. Gibbs, Jr., and Victor 
G. Springer of the U.S. National Museum. 

The work was done at the Department of Biological Sciences, 
Division of Systematic Biology of Stanford University, at the Depart- 
ment of Anatomy, Stanford University School of Medicine, and at 
the U.S. National Museum, Smithsonian Institution. 



SOUTH AlIERICAN CHARACID FISHES — ^WEITZMAN 129 

Material Examined 

This study is based on the examination of the skeletal preparations 
listed below. All specimens are alizarin preparations unless otherwise 
noted. In addition, considerable reference has been made to osteo- 
logical preparations listed in Weitzman (1962, pp. 11-17). The 
methods of preparing specimens for osteological investigation are the 
same as those I used earlier (Weitzman, 1954; 1962). 

The osteological drawings of Poecilobrycon harrisoni Eigenmann are 
based on one specimen (Stanford University 50245), 36.4 mm. in 
standard length. This specimen, together with eleven others, was 
found in the collections of the Division of Systematic Biology of 
Stanford University. These specimens had no data other than 
"British Guiana, Georgetown." Six additional aquarium-reared 
specimens (Stanford University 50244), 9.5 to 21.0 mm. in standard 
length, were stained with alizarin and compared with the specimen 
used for drawing. Since general proportions and shapes in the osteo- 
logical drawings were dehneated by optical methods, some distortion 
can be found in the figures. This is especially noticeable in figures 2-6, 
wherein the anterior cranial and snout region is somewhat fore- 
shortened. All illustrations are by the author. 

In the Ust below, CAS refers to the California Academy of Sciences; 
CAS(IUM) refers to specimens belonging to the CaUfornia Academy 
of Sciences but previously deposited at Indiana University and still 
bearing an Indiana University number; SU refers to specimens be- 
longing to the Division of Systematic Biology, Department of Bio- 
logical Sciences, Stanford University; USNM refers to the United 
States National Museum; ANSP refers to the Academy of Natural 
Sciences of Philadelphia. 

Copeina guttata (Steindachner) : SU 51692, two (of four), SL 56.5-57.0 mm., 
Peru, creek near Yurimaguas, November 1920, W. R. Allen. 

Copella nattereri (Steindachner): CAS 20743, four (of 35), SL 23.5-35.5 mm., 
Brazil, State of Pard, Lagoa Grande [probably Lagoa Grande do Javari about 
35 mi. northwest of Santar^m] July 17, 1924, Carl Ternetz. 

Erythrinus erythrinus (Bloch and Schneider) : SU 57678, four, SL 52.0-99.5 mm., 
Peru, creek near Yurimaguas, November 1920, W. R. Allen. 

Hoplerythrinus unitaeniatus (Agassiz) : CAS(IU]VI) 17106, one (of four) , SL 97.0 mm. 
Bolivia, Cuchuela [Cachuela] Esperanza, March 1922, N. E. Pearson; USNM 
163188, one (of four), SL 86.9 mm., Venezuela, a few miles off Puerto Aya- 
cucho, "Amazon territory," March 13, 1950, J. A. Rivero. 

Hoplias malabaricus Bloch: CAS(IUM) 17107, five, SL 31.0-108.0 mm., Bolivia, 
Huachi, at junction of the Ri'o Bopi and Rio Cochabamba [Santa Elena], Rio 
Beni basin, September-October 1921, N. E. Pearson; SU 3106, one, dry skele- 
ton, cranium 31 mm. in total length, Brazil, Maraj6 Island, Rio Tocantins, 
C. F. Hart. 



130 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

Lebiasina bimaculata Valenciennes: CAS(IUM) 15171, three (of 30), SL 57.5- 
60.9 mm., Peru, Piura, January 11, 1919, Carl H. Eigenmann. 

Nannostomus beckfordi Giinther: SU 50258, six (of 44), SL 22.0-26.5 mm., origi- 
nally from SU 50257, Brazil, State of Pard, south bank of Rio Amazonas, 
"Rfo Urara," June 26, 1924, Carl Ternetz. 

Nannostomus digrammus Fowler: SU 50249, two, SL 16.3-19.4 mm., Brazil, 
State of Amazonas, Manaos [Manaus], 1865, Louis Agassiz; SU 50251, three, 
SL 18.0-22.8 mm., Brazil, State of Amazonas, Igarapd do Mai Joana, a tribu- 
tary of the Rio Negro near Manaos [Manaus], December 25, 1924, Carl Ternetz. 

Nannostomus espei (Meinken) : ANSP 73873, one, SL?, British Guiana, an un- 
named creek, tributary to the Paruma River, itself tributary to the Mazaruni 
River via the Kamarang River, Pakaraima Mountain region of western British 
Guiana, 1955, Louis Chung; SU 50252, one, SL 25.0 mm., aquarium specimen, 
locality data unknown but probably the same as ANSP 73873. 

Nannostomus marginatus Eigenmann: SU 54119, three (of 32), SL 19.0-21.5 mm., 
originally from SU 50219, Brazil, State of Pard, Lagoa Grande [probably Lagoa 
Grande do Javari about 34 mi. northwest of Santar^m], August 20, 1929, Carl 
Ternetz. 

Nannostomus trifasciatus Steindachner: SU 50223, one, SL 29.5 mm., British 
Guiana, no other data; SU 54130, one (of two), SL 31.5 mm., aquarium speci- 
men, "from the Amazon," sent to G. S. Myers by Frederick Stoye. 

Piabucina festae Boulenger: SU 51068, four (of 12), SL 60.5-71.5 mm., Colombia, 
Rio Truand6, a western tributary of the Ri'o Atrato near Rio Sucio, 1913, 
C. E. Wilson; USNM 167795, one (of six), SL 70.0 mm., same data as preceding. 

Piabucina panamensis Gill: USNM 109234, two (of seven), SL 77.7-84.0 mm., 
Panama, Cativd,, March 2, 1937, S. F. Hildebrand. 

Piabucina erythrinoides Valenciennes: USNM 121400, three (of 27), SL 35.1-101 
mm., Rio Chama at Estanques, State of M^rida, Venezuela, April 3, 1942, 
L. P. Schultz. 

Piabucina species?: USNM 123796, two (of six), SL 64.1-67.3 mm., Colombia, 
Rio Magdalena, Cecil Miles. 

Poecilobrycon eques (Steindachner): SU 50247, two (of five), SL 28.5-30.0 mm., 
originally from SU 50229, Brazil, State of Amazonas, Sao Gabriel [Uaupes], 
Rio Negro, rockpools below rapids, February 1, 1925, Carl Ternetz; SU 50224, 
three, SL 25.8-32.5 mm., aquarium specimens, no other data. 

Poecilobrycon harrisoni Eigenmann: SU 50246, one, SL 29.5 mm., originally from 
SU 50245, British Guiana, Georgetown, collector unknown; SU 50245, one 
(10 unstained), SL 36.4 mm., British Guiana, Georgetown; SU 50244, six, 
SL 9.5-21.0 mm., aquarium specimens, reared by S. Weitzman. 

Poecilobrycon unifasciatus (Steindachner): CAS(IUM) 11704, one (of 10), SL 
33.0 mm., British Guiana, Rockstone sandbank, Essequibo River, 1908, 
C. H. Eigenmann; SU 50268, two, SL 27.8-30.4 mm., aquarium specimens, 
no other data. 

Pyrrhulina filamentosa Valenciennes: CAS(IUM) 12186, four, SL 14.7-20.2 mm., 
British Guiana, Aruka River, 1908, C. H. Eigenmann. 

Pyrrhulina semifasciata Steindachner: CAS(IUM) 12172, one, SL 58.1 mm., 
British Guiana, Holmia Creek, 1908, C, H. Eigenmann. 

Pyrrhulina spilota Weitzman: USNM 197523, two (of 19), SL 41.5-50.4 mm., 
first generation descendants of holotype and paratype. 

Pyrrhulina vittata Regan: USNM 197524, SL 26.0 mm., aquarium specimen, 
locality unknown. 



SOUTH AMERICAN CPIARACID FISHES — WEITZMAN 131 

Historical Review of the Classification of Nannostomus and 

Poecilobrycon 

Giinther (1872, p. 146) described the first member of the Nannos- 
tomina, Nannostomus beckfordi. Of its relationships with other 
characids, he simply stated: ". . . allied to Lebiasina, but with a 
totally different form of the snout and mouth." He considered it to 
belong to his first group of characids, the Erythrina (Giinther 1864, 
pp. 278 and 281), which included the genera Hoplias, Erythrinus, 
Lebiasina, Pyrrhulina, and Corynopoma. They were defined as those 
characids without an adipose fin. 

Steindachner (1876, p. 130) thought Nannostomus should be placed 
in or near the group Anostomatina ^ of Giinther (1864, pp. 279, 303). 
Giinther's Anostomatina included the genera Anostomus, Rhytiodus, 
and Leporinus. Steindachner (1876, p. 122) pointed out that the 
presence or absence of the adipose fin in Nannostomus (=Nannos- 
tomus plus Poecilobrycon of later authors) is not of importance in 
determining their relationships with other characids because this fin 
may be present or absent in specimens of Poecilobrycon eques. 

Eigenmann and Eigenmann (1891, p. 49) placed Nannostomus 
in their subfamily Anostomatinae, which also included Anostomus, 
Laemolyta, Characidium, Rhytiodus, Leporellus, and Leporinus. 
Boulenger (1904, p. 576), following Steindachner 's and Eigenmann 's 
precedent, placed the genus Nannostomus in the subfamily 
Anostominae. 

Eigenmann (1909b, p. 35-36) placed the genera Nannostomus, 
Poecilobrycon, and Archicheir (the latter two therein described) in his 
undefined subfamily Nannostomatinae; however, the name Nan- 
nostomatinae first appeared in Eigenmann (1909a). He also included 
the genera Characidium and Microcharax in this subfamily. Eigen- 
mann (1910, p. 427) again listed his subfamily Nannostomatinae 
and, in addition, included the genus Jobertina. Eigenmann may have 
held the opinion that the members of his subfamily Pyrrhulininae 
are aUied to Nannostomus and Poecilobrycon for he placed them 
immediately after his Nannostomatinae. These two subfamilies 
were separated widely from his Erythrininae. Eigenmann's Anosto- 
matinae (including Leporinus and relatives) immediately preceded 
his Nannostomatinae. Apparently at that time he thought them 
related. Eigenmann (1912, p. 254), in a key to the subfamilies 
of characids from British Guiana, first defined the subfamily Nan- 



2 According to Miller (1897, p. 132), family group names formed from adjec- 
tives used substantively, e.g., Nannostomus and Anostomus, should not be ter- 
minated by "-atidae" or "-atinae." Accordingly, Giinther's Anostomus should 
become Anostomina, not Anostomatina, or Anostominae, not Anostomatinae. 



132 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

nostomatinae. There are three important errors in Eigenmann's 
concept of this subfamily's morphology. Teeth are present in the 
maxillary, not absent; the gill membranes are joined to each other 
but free from the isthmus, not slightly united; and the parietal 
fontanel is absent, not present as Eigenmann indicated. 

Regan (1911, p. 21) placed the genera Nannostomus and Characidium 
in the Nannostominae and included this subfamily in his family 
Hemiodontidae. This family consisted only of his Hemidontinae 
and Nannostominae. 

Cockerell (1914, p. 98-99), in a study of characid scales, noted the 
close resemblance of the scales of Nannostomus and Pyrrhulina and, 
on this basis alone, found reason to more closely associate Pyrrhulina 
with Nannostomus than Poecilobrycon with Nannostomus. He also 
noted the very different structure of the scales of Characidium. I 
find that the scales of Poecilobrycon and Nannostomus are much alike 
and, in addition, are very similar to those of Pyrrhulina, Copella, 
and Copeina. In confirmation of part of CockerelPs work, however, 
the osteology and scales of CJiaracidium differ quite widely from the 
osteology and scales of any of the other genera discussed here. 

Gregory and Conrad (1938, pp. 324, 344-347) followed Regan 
(1911) in relating Nannostomus Si.nd Poecilobrycon to Hemiodus smd 
its relatives. They placed Nannostomus and Poecilobrycon in a sub- 
family (Hemiodontinae) considered by them to be widely separated 
from the subfamily (Characinae) containing Pyrrhulina. As shown 
below, my work indicates that Nannostomus, Poecilobrycon, and 
Pyrrhulina are related rather closely and are placed in the tribe 
Pyrrhulinini. 

Fowler (1950, pp. 253-263) considered the genera Characidium, 
Microcharax, Nannostomus, Archicheir, and Poecilobrycon as con- 
stituting the subfamily Nannostominae (apparently following Eigen- 
mann, he spelled it Nannostomatinae) . He placed this subfamily 
nearest his subfamilies Leporininae and Parodontinae but did not 
comment on relationships or define his groups. 

Hoedeman (1950a, p. 14) established the tribe Nannostomini to 
include the genera Nannostomus, Poecilobrycon, and his newly pro- 
posed N annobrycon. He apparently did not consider that the genus 
Archicheir belonged to his Nannostomini, for he excluded it from his 
treatment. 

In his first paper on Nannostomus and relatives, Hoedeman (1950a, 
p. 11) considered his Nannostomini to be related to the Hemiodon- 
tinae; however, he excluded Characidium from relationship with the 
Hemiodontinae. He presented no evidence for this opinion. Hoede- 
man (1954a, p. 83) reconsidered his classification and placed the tribe 
(emended to Nannostomidi) in his family Erythrinidae, a family he 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 133 

then considered as comprising two subfamilies, Erythrininae and 
Anostominae. He here excluded the Hemiodontinae from his Ery- 
thrinidae. His subfamily Erythrininae comprised four tribes, Lebia- 
sinidi, Erythrinidi, FyrrhuHnidi, and Nannostomidi. He did not 
present evidence for this classification but he did remark (words in 
brackets are mine) : "Poecilobrycon cannot be derived from either of 
these genera [i.e., Nannostomus or Nannobrycon], nor vice versa. The 
two groups probably had a common ancestor, close to the present day 
Pyrrhulina, to which both are more closely related than to each other." 
Hoedeman, however, placed PoecUobrycon, Nannostomus, and Nanno- 
brycon in one group and the Pyrrhulina in another. As will be shown 
below, it is my opinion that the members of the Nannostomina as 
defined below are more closely related to each other than to any other 
characids, but that Hoedeman was correct at that time in suspecting 
their relationship to be with Pyrrhulina. Hoedeman (1954b, pp. 68- 
84, and 1956b, pp. 547-551), however, once again reconsidered his 
classification of Nannostomus and PoecUobrycon and placed them with 
Hemiodus and relatives, excluding them from close realtionship with 
Pyrrhulina. 

In summary, the Nannostomina have been thought to be related 
to Lebiasina and its relatives, Erythrinus and its relatives, Pyrrhulina 
and its relatives, Characidium, and finally to Hemiodus and its rela- 
tives. As will be shown, their morphological relationships are with 
Pyrrhulina, Lebiasina, and close relatives. Inadequate morphological 
investigations have been the primary cause of this divergence of 
opinion. 

Historical Review of the Classification of the Erythriainae 
and Lebiasininae 

Since in the present work Nannostomus and PoecUobrycon have 
been found to belong to the subfamily Lebiasininae, a historical 
review of their relationships to other characids would not be complete 
without a historical review of the Lebiasininae. Because the Lebia- 
sininae and Erythrininae have been associated closely by many 
authors, both are reviewed. 

Valenciennes, in Cuvier and Valenciennes (1846, p. 480), estab- 
lished the "famille des Erythroides" for the reception of the genera 
Macrodon (=Hoplias), Erythrinus, Lebiasina, and Pyrrhulina. He 
defined these fishes as having a double swim bladder that is some- 
times cellular, teeth on their jaws and palate, the cheek covered by a 
large subopercle, the belly always rounded, and no pyloric caeca. 
Apparently, Valenciennes interpreted the small autogenous "supra- 
opercular" element of Hoplias as the opercle and the large true opercle 
as the subopercle. 



134 PROCEEDINGS OF THE NATIONAL MUSEUM ^"^- ^^^ 

Giinther (1864, pp. 278, 281-288) added the genus Corynopoma 
to Valenciennes' family grouping. This genus originally was con- 
sidered by Gill (1958, pp. 422-428) to have affinities with the Chara- 
cinae. Regan (1906, p. 382; 1911, p. 16) has shown good reasons for 
not placing Corynopoma with the genera Hoplias, Erythrinus, and 
Hoplerythrinus . 

Gill (1858, pp. 410-413) estabhshed the family Erythrinidae for 
Pyrrhvlina, Erythrinus, and Lebiasina. Eigenmann and Eigenmann 
(1889, pp. 100-115) followed Giinther (1864) in including Corynopoma 
in what otherwise essentially equals GiU's family Erythrinidae. 
Neither Giinther nor Eigenmann and Eigenmann had seen examples 
of GiU's Corynopoma. The Eigenmanns, however, chose to give the 
group subfamily rank (Erythrininae) within the Characidae. The 
Eigenmanns listed characters for the Erythrininae as foUows: 

Adipose fin none. Gill opening wide, the membranes slightly united, free 
from the isthmus. Nares approximated. Teeth well developed, at least 
in the jaws; pharyngeal teeth villiform. Cheeks covered by the suborbital 
bones. Brain case entirely closed above. Body elongate, slender, fusiform 
or subfusiform. Back not greatly arched, belly rounded. Dorsal short, of 
8 to 15 rays. Intestine short. Carnivorous. 

This definition does not exclude aU members of the Characinae or 
members of some of the other subfamiUes of characids. 

Regan (1911) included all the members of Gill's Erythrinidae 
in his Characidae; however, he separated Pyrrhvlina widely from 
Lebiasina and Piabucina because he found that Pyrrhulina lacked 
ectopterygoid teeth while Lebiasina and Piabucina possessed them. 
Examination of many characids indicates that the presence or absence 
of ectopterygoid teeth is of minor importance because, in some species 
that normally possess them, they may be unilaterally or bilaterally 
absent in a few specimens. They may also be present in one species 
but absent in otherwise closely related species, e.g., Charax and its 
relatives.' 

Regan (1911) maintained Erythrinus, Hoplerythrinus, and Hoplias 
in the subfamily Erythrininae and the genera Lebiasina and Piabucina 
in the subfamily Lebiasininae. He placed Nannostomus and Poecilo- 
brycon in his Hemiodontidae because their premaxillaries are movable, 
because he thought they had no ectopterygoid teeth, and because he 
misinterpreted the palatine arch. 



* The establishment of the family Acestrorhynchidae by Fernandez- Yepez 
(1955, p. 450) and the genus Charaxodon by Fernandez- Yepez (1947, p. 1), based 
principally, if not whoUj^, on the presence or absence of ectopterygoid teeth 
(= the teeth of the "palatine" of Fernandez- Yepez) should be reviewed with a 
more critical approach. Bohlke (1958, p. 70) considered Charaxodon to be a 
synonym of Morallesia. 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 135 

Regan (1912) lumped the genera Chalceus, Pyrrhidina, Copeina, 
and Pogonocharax into a "natural group," stating that they differed 
from the rest of the Characidae by a very large mesethmoid (= ethmoid 
of this work), oblong or elongate body form, rounded abdomen, 
fiattish upper surface of the head, large scales, and short dorsal 
and anal fins. These are superficial characters and do not indicate 
true relationships. The osteology of Chalceus appears to be in many 
respects like that of Brycon and there is reason to beheve that 
Chalceus may have been derived from Brycon or a very close ancestor. 
In any event, its osteology is typically that of the Characinae as 
defined by Weitzman (1962). The case of Pogonocharax rehi, a fish 
described by Regan and presumed by him to be from South America, 
was reviewed by Myers (1956b, p. 13); the fish is not a characid but 
an Asiatic cyprinid of the genus Esomus. 

Gregory and Conrad (1938, pp. 324, 343-344) recognized the sub- 
family Erythrininae for the inclusion of the Erythrininae and Lebi- 
asininae of Regan (1911). Superficially, Regan's two subfamilies do 
look much alike but they are separated by the characters listed below 
in the classification. Gregory and Conrad (1938, p. 343) have noted 
the resemblance of Hepsetus (= Sarcodaces) to Hoplias; indeed, 
Hepsetus possesses a supraopercular like Hoplias. The cranial bones 
do have a superficial and perhaps even a phylogenetically significant 
resemblance to those of Hoplias. Basically, however, the osteology 
of Hepsetus is more like that of the Characinae, especially with regard 
to the pectoral girdle. In addition, Hepsetus has four branchiostegal 
rays, a clear anastomosis between the dermopterotic and supraorbital 
laterosensory canals, no accessory ectopterygoid, and the usual 
characid number of eight orbital bones. Certain features of the skull, 
however, such as the frontal-sphenotic articulation and the presence 
of a supraopercular are characters in common with the Erythrininae but 
not the Characininae and suggest that'" the relationships of Hepsetus 
should be investigated further. In addition, the possible relationships 
of Ctenolucius and Boulengerella with Hepsetus should not be ne- 
glected. Preliminary examination indicates that the relationships of 
Acestrorhynchus and Acestrorhamphus are probably with Charax, 
Roeboides, and other closely related genera in the Characinae, not with 
Boulengerella and Ctenolucius as assumed by Gregory and Conrad 
(1938, pp. 323-324, 338-344). These authors followed Regan (1911) 
in placing Nannostomwi and Poecilobrycon with members of the 
Hemiodontinae. 

Hoedeman (1954b, p. 55 ; 1956a, p. 12) presented a classification of the 
major characid groups. Most of his work appears to be speculation. 
The only morphological data of any value was a superficial comparison 
of scales from several characids. In his 1956 classification, Hoedeman 



I 



136 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

separated charaeids into three families, one of which was the Eryth- 
rinidae. He divided this family into two subfamihes. The first, the 
Erythrininae, contains his Lebiasinidi (= Lebiasinini of the present 
work), Erythrinidi (= Erythrininae), Pyrrhulinidi (= Pyrrhuhnina) 
and Nannostomidi (== Nannostomina). The second subfamily of 
Hoedeman's Erythrinidae is the Anostominae. In this group 
he placed four tribes, Curimatidi, Anostomidi, Hemiodontidi, and 
Prochilodidi. Hoedeman (1956a, p. 12) stated: 

Vers le milieu de I'eocene, les Erythrinidae se divisent en deux sous-families: 
(a) Erythrininae, avec les tribus actuelles: (1) Lebiasinidi, (2) Erythrinidi, 
(3) Pyrrhulinidi, et (4) Nannostomidi; et la seconde sous-families: (b) 
Anostominae, avec les tribus: (5) Curimatidi, (6) Anostomidi, (7) Hemio- 
dontidi, et (8) Prochilodidi. 

There is no fossil evidence for this statement nor is there fossil evi- 
dence for any of the other of Hoedeman's phylogenetic and time- 
sequence inferences and speculations given in his paper. 

Piton (1938) described two fossil fishes from the mid-Eocene at 
Menat, France. Piton thought that these fishes, Prohydrocyon pelle- 
grini and Procharacinus arverniensis, were fossil charaeids. Perhaps 
Hoedeman's use of the date of mid-Eocene is from Piton 's paper. It 
appears now that there is no evidence for charaeids from the mid- 
Eocene of France (Weitzman, 1960). 

The original evidence presented by Hoedeman (1956a) for his phy- 
logeny was apparently derived from examination of a few representa- 
tive characid scales. Examination of characid scales of the genera 
listed in the material section of Weitzman (1962) shows a greater 
variation in the scale morphology of characid groups than Hoedeman 
found. Sufficient variation was found among the representatives of 
the various groups to indicate that, although scales wiU prove useful 
as an aid in establishing a classification and for drawing phylogenetic 
inferences, their use alone, without an attempt to correlate them with 
many other morphological characters, may more often prove mis- 
leading than helpful. 

Osteology of Poecilobrycon harrisoni and Related Charaeids 

In the osteological discussion below, parts of the skeleton of Poecilo- 
brycon harrisoni are described and compared with other species of 
Poecilobrycon and Nannostomiis. Also, other characid genera and 
groups thought, either by the present author or other authors, to be 
pertinent to a discussion of the relationships of the Nannostomina 
are compared with the basic skeletal plan of Nannostomiis and 
Poecilobrycon. 

Cranium (figs. 2, 3, 4, 5, 6). — The median ethmoid bone is a large, 
rather thin plate that has a profile in dorsal view as shown in figure 2. 



SOUTH AMERICAN CHAR ACID FISHES — WEITZMAN 137 

This bone has bilateral posterior lamellae projecting backward under 
the frontals and contacting the dorsoanterior portion of each lateral 
ethmoid. The ethmoid probably is mostly of supraethmoidal (der- 
methmoidal) origin, but an anterior, ventral lamella of the ethmoid 
that extends inferiorly and posteriorly to contact the prevomer below 
is probably of endochondral origin. The ethmoidal spine found in 
most characids (see Weitzman, 1962, figs. 2, 3, 4 of Brycon meeki) is 
reduced to a very small structure. The premaxillaries are not firmly 
attached (the joint being a loose syndesmosis) and are somewhat 
movable, being attached to the ethmoid by fairly long, lax ligamentous 
tissue. They cannot, however, be described as protractile. The 
cartilagenous part of the ethmoid is restricted to the area between 
the large prevomer and the ethmoid. A few foramina are present on 
the dorsoanterior surface of the ethmoid. All species of Nannostomus 
and Poecilobrycon have a very similarly shaped ethmoid; however, 
members of the genus Nannostomus have a shorter snout and, there- 
fore, a shorter, more blunt ethmoid than members of the genus 
Poecilobrycon. 

The large ethmoid of the genera Pyrrhulina, Copella, and Copeina 
is very similar to that in the Nannostomina in its relationships to the 
prevomer, lateral ethmoids, and frontals. It differs, however, in 
having a broader, more rounded horizontal profile and a very well- 
developed, though rather obtuse, ethmoidal spine. In the Pyrrhuli- 
nina the premaxillaries are attached to the ethmoid by a very slightly 
movable syndesmotic fibrous joint. The ethmoid of Lebiasina and 
Piabucina is generally similar in structure to that of Pyrrhulina, 
Copella, and Copeina, but proportionately the ethmoidal spine is 
considerably smaller. 

The ethmoids of the Lebiasinini and Pyrrhulinina differ most promi- 
nently from those of the Nannostomina in the following manner: The 
ethmoidal spine is well developed, the premaxillary is firmly fixed to 
a groove along the anterolateral border of the ethmoid by a tight 
fibrous joint, and the ethmoid bone is not as oblong or square in 
horizontal profile, being somewhat more wedge-shaped. 

The ethmoid of Hoplias (Starks, 1926, p. 160, fig. 8) and Erythrinus 
is quite similar in general form to that of Brycon (Weitzman, 1962); 
however, the joint between the ethmoid and prevomer is bordered by 
a lateral lamina of cartilage that is in contact with, or approaches, a 
mass of cartilage on the medial anterior border of the palatine (fig. 
4). In the Characinae, only very small amounts of cartilage exist in 
these areas. The ventral diverging bony ethmoid lamellae, so well 
developed in the other characid fishes examined, are not at all, or only 
weakly, developed in the Nannostomina, only weakly so in the Pyr- 
rhulinina and the Lebiasinini, but well developed in the Erythrininae. 



138 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

The toothless prevomer is concave ventrally and concave dorso- 
laterally, being an inverted Y-shaped bone in cross section. In small 
specimens of Poecilobrycon harrisoni and in the large adults of some 
species (Nannostomus beckfordi and N. trifasciatus) , the prevomer is 
formed of three thin concave plates of bone adhering to, or about, 
a central cartilaginous core. The two fused dorsal plates contact the 
ethmoid by a cartilaginous (synchondral) joint and posteriorly by 
another synchondral joint to the parasphenoid and lateral ethmoid. 
There is no rhinosphenoid. The posterior shaft of the prevomer, 
which ordinarily contacts the parasphenoid in the characinae, is ex- 
tremely short and does not reach the parasphenoid. There are a 
number of foramina on the ventral surface of the prevomer. Many of 
these are probably for branches of the ramus buccalis facialis nerve. 

Members of the Erythrininae, Lebiasinini, and Pyrrhulinina have 
a prevomer essentially like that of the Nannostomina; however, the 
shaft of the vomer is better developed in groups other than the Nan- 
nostomina and extends well back onto the parasphenoid. Except for 
the reduced prevomerine shaft in the Nannostomina, the prevomer of 
the Lebiasininae is essentially like that of the Characinae. 

Each lateral ethmoid projects downward from under its respective 
frontal and contacts its counterpart at the median vertical plane by 
a nonmovable synchondral (cartilaginous) joint. The foramen for 
the olfactory nerve is near the median edge of the lateral ethmoid. 
An upper medial blade of the lateral ethmoid extends anteriorly and 
medially to contact a vertical median wall of cartilage behind the 
prevomer and there forms a nonmovable synchondral joint. Antero- 
ventrally the lateral ethmoid has a process extending forward to contact 
a sheet of cartilage that extends anteriorly to the lower portion of 
the prevomer. Ventrally the prevomer is in contact through cartilage 
with the parasphenoid and posteromedially with the orbitosphenoid. 
The lateral ethmoid of all members of the Nannostomina and other 
members of the Lebiasininae is essentially the same. In the Pyr- 
rhulinina and the Lebiasinini the distance between the prevomer and 
the lower medial portion of the lateral ethmoid is much shorter. That 
of young specimens of Hoplias is similar to that in the Nannostomina, 
but in older specimens the two bones meet each other. The structural 
configuration of the lateral ethmoid in the Erythrininae and Chara- 
cinae is not markedly different in the two groups, and those differences 
in form that do occur can be correlated with differences in the shape 
of mouth and snout and their functions. 

The frontals of Poecilobrycon harrisoni are large, smooth, gently 
curved bones. They are similar to the frontals of the Characinae in 
their basic structm-al relationship to other bones of the cranium; 
however, the frontal-parietal fontanel is never present in adults and 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 139 

the epiphyseal bar is relatively small in size. lu young specimens a 
well-developed fontanel is present. A specimen 14.8 mm. in standard 
length had a well-developed fontanel while a specimen 21.8 mm. in 
standard length had none. The supraorbital laterosensory canal in 
the frontal has the normal characid com'se but with the following 
exceptions : The portion of the canal that normally is buried in frontal 
bone over the epiphyseal bar (the mesial or epiphyseal branch) is 
reduced greatly in length, and the canal that in the Characinae passes 
backward in the frontal to the pareital (the posterior branch) is 
absent. 

The distribution of major laterosensory canals is the same in the 
frontal of the Pyrrhulinina as in the Nannostomina although the 
mesial branch (supraorbital commissure of Branson and Moore, 1962) 
sometimes is developed better in the Pyrrhulinina. In Piabucina the 
posterior branch is present but it does not reach the parietal. In 
both Lehiasina and Piabucina there is a branch of the dermopterotic 
sensory canal extending backward in the lateralmost edge of the 
frontal. Posteriorly the dermopterotic canal enters the pterotic. 
Anteriorly this canal enters the infraorbital canal. In Hoplias, 
Erylhrinus, and Hoplerythrinus the lateral edge of the frontal bone 
extends outward to contact the spinous process of the sphenotic, 
completely covering and encircling the dilator groove. This is not 
true in the Characinae, nor in the Lebiasinini, Pyrrhulinina, or 
Nannostomina. In these fishes the dilator groove is covered but not 
encircled by the frontal. Small specimens of Hoplias have the 
frontals and parietals separated by a fontanel, whUe adults do not. 
In Hoplias and Erylhrinus the supraorbital sensory canal is the same 
as in the Nannostomina, but the posterior branch extends into the 
parietal. 

The supraoccipital of Poecilobrycon harrisoni is not unlike that of 
Brycon in its position and relationships to other cranial bones. The 
supraoccipital of the Pyrrhulinina, Lebiasinini, and Erythrininae is 
quite similar to that of the Nannostomina and differs mainly from the 
Characinae in the replacement of the supraoccipital spine with a 
crest. 

The exoccipitals, basioccipital, and epiotics of Poecilobrycon har- 
risoni are very much like those of the members of the Characinae and 
differ in a manner correlated with the widening and flattening of the 
skull. This may best be seen by comparing the illustration in 
Weitzman (1962, fig. 5) with figure 5 of the present work. The 
myodome is much reduced in its vertical depth. The exoccipitals, 
epiotics, and basioccipital of the Pyrrhulinina, Lebiasinini, and 
Erythrininae differ from those of the Characinae in having shapes 
similar to those of the Nannostomina. 



140 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

The opisthotic is a small tabular boue applied to the joiut between 
the pterotic and exoccipital but lying mostly on the pterotic. 

Dorsally the pterotic is overlaid by the parietal and posteriori}'- 
contacts the epiotic. Anteriorly and dorsally the pterotic does not 
reach the frontal but is completely bounded by the sphenotic, while 
medially it contacts the prootic. Except for lacking a dermopterotic 
laterosensory canal, for not contacting the frontal, and except for its 
general shape, the pterotic is much like that of the Characinae. The 
pterotic of the Pyrrhulinina is like that of the Nannostomina in 
lacking a laterosensory canal and in not being jointed with the frontal. 
The Lebiasinini and the Erythrininae have a laterosensory canal in 
the pterotic and the frontal contacts the sphenotic. 

The sphenotics are much like those of the Characinae; their various 
relationships to other bones have been described under the frontals 
and parietals. Their relationships to the semicircular canals are 
the same as in the Characinae. 

In their essential relationships to other bones and the soft anatomy, 
the prootics are not greatly different from those of the Characinae. 
Detailed comparison of the various foramina of this bone with the 
foramina of the prootics of the Pyrrhulinini, Lebiasinini, and Ery- 
thrininae undoubtedly would prove productive in determining rela- 
tionships; however, this work must be postponed pending detailed 
studies of the nerves, veins, and arteries passing through this com- 
plicated bone. 

The subtemporal fossa is present as a shallow groove along the 
joint between the prootic and pterotic and is present in all members 
of the Erythrininae and Lebiasininae. 

The pterosphenoid of Poecilobrycon harrisoni and other species of 
the Nannostomina is similar to that in the Characinae, being a large 
plate in the upper posterior wall of the orbit. The foramen for the 
trochlear nerve is located at the junction of the articulation between 
the frontal, pterosphenoid, and orbitosphenoid. 

The orbitosphenoid is Y-shaped in cross section; the upper arms 
of the Y are jointed to the frontals. A foramen for the first cranial 
nerve is enclosed partially by the orbitosphenoid. Other species in 
the Nannostomina examined also have an orbitosphenoid that is 
Y-shaped in cross section. The same is true of the Pyrrhulinina. 
In the Lebiasinini the cross sectional shape of the orbitosphenoid is 
more like a V than a Y. In Hoplias the foramen for the first cranial 
nerve is included completely in the orbitosphenoid of adults but 
included only partially in that of the young. Erythrinus has a rather 
peculiar orbitosphenoid, consisting of a pair of bilateral flat plates 
attached ventrally by cartilage. In specimens larger than those 
examined these plates possibly may fuse ventrally, producing a more 



SOUTH AJVIERICAN CHARACID FISHES — WEITZMAN 141 

"normal" orbitospheiioid. The orbitospheuoid does uot have a 
foramen for the fu"st cranial nerve in the small specimens of the 
Erythrininae examined, but it may, in larger specimens. An X-ray 
negative of a specimen of Hoplias, 108 mm. in standard length, 
indicates that a foramen may be included partially in the orbitospheuoid. 

Orbital bones (fig. 6). — There are six infraorbital bones in the 
Lcbiasininae and Erythrininae. The supraorbital is absent. The 
first and second infraorbitals in adult specimens of Poecilobrycon 
harrisoni have a laterosensory tube imbedded within them. The 
remainder of the infraorbital bones have the infraorbital canal along 
their ocular edge but it is enveloped only partially in bone. Only 
Poecilobrycon eques, P. harrisoni, and P. unifasciatus have a sensory 
tube contained within the bony substance of both the first and second 
infraorbitals. All other Imown members of the Nannostomina lack 
this bony tube in the second infraorbital, the infraorbital canal 
passing over the surface of the bone. The antorbital is present and 
weU developed in the Nannostomina; it sometimes partially envelops 
the anterior part of the infraorbital canal. In the Pyrrhulinina the 
first two infraorbital bones have enclosed canals; sometimes the 
other infraorbital bones also have enclosed canals in members of 
this subtribe. 

In the Lebiasinini the infraorbital canal may be included within 
the bony substance of all the orbital bones or it may pass only over 
the surface of the second through the fifth infraorbital. It may pass 
either within or over the substance of the antorbital. 

In the Erythrininae the orbital bones have an infraorbital canal 
imbedded in them. There is no supraorbital; the antorbital contains 
a canal and is fused with the first infraorbital. This is apparently 
similar to the situation found in certain cyprinodonts and catfishes 
(see Weitzman, 1962, pp. 28-31). Another possible interpretation is 
that the antorbital is absent, the first infraorbital having extended 
into the topographical area of the antorbital. 

The elongate nasal bone of the Nannostomina has the usual char- 
acid relationship, being a tubular bone connected with, and anterior 
to, the supraorbital canal of the frontal. In the Pyrrhulinina, Lebias- 
inini, and the Erythrininae the nasal bone is not greatly different 
from that of the Nannostomina except for differences correlated with 
the broader and shorter snouts in these fishes. 

Opercular apparatus (figs. 6, 7). — The opercular bones of the 
Nannostomina differ from those of the Characinae mainly in having 
a different shape correlated with the elongate and relatively flattened 
head. The thin interopercle internally covers much of the lower sur- 
face of the symplectic, preopercle, metapterygoid, and even some of 
the quadrate. The preopercle does not extend forward to near the 



142 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

mandible as it does in the Characinae, Pyrrhulinini, Lebiasinini, and 
Erythrininae, but it is limited in its anterior extent to an area under 
the long, slender, lower, posterior process of the quadrate. The 
opercle and subopercle are similar in Lebiasininae and Erythrininae 
and are not markedly different from those of the Characinae. 

Upper jaw (figs. 6, 7). — The premaxillaries of the Nannostomina 
bear a single row of one to seven small cuspid teeth. A row of "re- 
placement" teeth is present behind the main outer row. The pre- 
maxillary is not divided into upper and lower processes but is more or 
less lenticular in its horizontal outline. Its dorsal surface has a de- 
pression to receive a lateral process of the ethmoid bone. 

The maxillaries of all members of the Nannostomina are essentially 
alike. The maxillary has a single tooth with from one to six cusps. 
The maxillary is a flat lamella of bone with a robust, conic upper 
process extending toward the anterior part of the ethmoid. Pos- 
teriorly the maxillary does not reach the second infraorbital. 

The premaxillary of the Pyrrhulinina bears one to two rows of 
simple conic teeth while the maxillary has a single row of conic teeth 
along its edge. The maxillary is elongate and reaches well along the 
second infraorbital to about the third infraorbital bone. The pre- 
maxillaries of members of the Pjrrrhulinina is not well divided into 
upper and lower processes. 

The premaxillary in members of the Lebiasinini has a single row of 
tricuspid teeth similar in general shape to those of the Nannostomina. 
The premaxillary is fairly well divided into an upper process and a 
lower lamella. The maxillary has a single row of peglike tricuspid 
teeth along its anterior edge, and its lower limb reaches to the third 
orbital bone. 

The premaxillary in members of the Erythrininae has well-developed 
upper and lower limbs and a single row of conical teeth. The max- 
illary is well developed and in Hoplias the fan-shaped distal end is 
external to the second infraorbital. In Erythrinus and Hoplery- 
thrinus the fan-shaped distal end of the maxillary is included under 
the second infraorbital. In the Erythrininae the maxillary almost 
reaches the third infraorbital. 

Lower jaw (figs. 6, 7). — The dentary of adult specimens of the 
Nannostomina bears two rows of teeth ; the anterior large teeth of the 
outer row have five or six cusps while the posteriormost few teeth may 
have the number of cusps reduced from one to fom*. The second row, 
composed of small conical teeth is on a crest of bone lying behind the 
replacement teeth of the first row. The shape of the dentary is rather 
remarkable and diagnostic of the Nannostomina. It is divided into 
two regions. The anterior region bears the teeth and is approxi- 
mately like the premaxillary in shape. The posterior portion is of 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 143 

the more normal characid shape and is of a thinner, lighter bone than 
the thick anterior region. The two regions are separated by a deep 
median groove and are joined by a latero ventral connecting sheet of 
bone. The complicated series of locking bony convolutions present 
at the symphysis of most characids is absent. The posterior portion 
of the dentary carries the tube for the laterosensory canal. 

The articular lies against the medial posterior surface of the dentary 
and bears a socket for the articular head of the quadrate. The 
angular is a well-developed bone ventral and internal to the lower 
posterior region of the articular and dentary. 

The coronomecklian (sesamoidarticular) is a prominent bone on 
the inside surface of the articular above the posterior region of Meckel's 
cartilage. 

The lower jaw in the Pyrrhulinina is similar in general form and 
shape to that of Brycon. The lower jaw of males of the genus Copella 
is somewhat modified and this is correlated with the differences 
found in the dentition and shape of the upper jaws (Myers, 1956a, 
p. 12). There are two rows of conical teeth on the dentary, an outer 
larger row and an inner row placed on a ridge of bone behind the 
replacement teeth of the outer row. There is a slight notch along 
the lower border of the dentary just posterior to the convoluted 
symphysis. This notch is probably homologous with the deep 
ventral groove in the dentary of the Nannostomina. The rest of 
the bones comprising the mandible in the Pyrrhulinina are similar 
in their basic relationships to those of the Characinae. 

In general shape the mandible in members of the Lebiasinini is 
like that of the Pyrrhulinina; however, the teeth in the outer row 
are tricuspid and similar in form to those found in the Nannostomina. 
The inner row is a series of very small conical teeth that, as in the 
subtribes Nannostomina and Pyrrhulinina, pass onto the lateral upper 
edge of the jaw behind the first large row of teeth. 

The lower jaw in the Erythrininae is not unlike that in the Lebia- 
sinini. In general, the placement of the dentary teeth in these two 
groups is much alike; however, in the Erythrininae the second row 
of teeth is restricted to the upper posterior edge of the dentary and 
to the area posterior to the first row of teeth. This second row does 
not advance forward on the crest of the bone behind the replacement 
teeth of the first and largest row of conical teeth. 

Mandibular and palatine arches (figs. 4, 6, 7). — Regan (1911, p. 
21) stated that Nannostomus and Characidium belong to the family 
Hemiodontidae and that these two genera could be distinguished 
from other hemiodontids by the possession of a single series of teeth 
in both jaws and a broad two-headed hyomandibular. As shown 
above, members of the Nannostomina have two rows of teeth in the 

728-42ft— 64 2 



144 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

lower jaw. Althougli examination shows that Characidium has a 
distinctly two-headed hyomandibular, that in the Nannostomina 
is single-headed. The hyomandibular in the Nannostomina is a 
broad bone with a broad dorsal articular surface that articulates 
with both the sphenotic and pterotic. It does not differ greatly in 
shape or relationships from the hyomandibular in the Characinae. 
The hyomandibular is not markedly different in shape or functional 
relationships in Pyrrhulinina, Lebiasinini, or Erythrininae. 

The quadrate of Nannostomus and Poecilobrycon is a large, rather 
rectangular bone with a long, slender posterior process below the 
symplectic. The dorsal surface of the quadrate extends upward, 
lateral to the large mesopterygoid, and the dorsal profile of the 
quadrate is gently curved. The posterior upper surface contacts 
the metapterygoid and anterior end of the symplectic thi-ough 
cartilage. The symplectic is quite large, elongate, and rather slab- 
shaped posteriorly. It is almost as long as the hyomandibular. 
The metapterygoid posteriorly contacts the lower end of the hyo- 
mandibular through cartilage. The central circular fenestra between 
the quadrate, symplectic, and metapterygoid, so typical of almost all 
other characids, is absent in the Nannostomina. Just anterior to 
the upper anterior corner of the quadrate is a smaU triangular ecto- 
pterygoid. In some members of the Nannostomina, this bone bears 
a few simple conical teeth. In one species, Poecilobrycon egues, 
teeth were foimd on the ectopterygoid of one side but not the other 
in some specimens. 

The toothless palatine is well developed and hes lateral to the 
pre vomer. The anterior head of the palatine Hes just posterior to 
the upper process of the maxiUary. 

The mandibular and palatine arches in the Pyrrhulinina and 
Lebiasinini examined are very similar to those in the Nannostomina, 
but, in general, these bones are not as elongate as in the Nannostomina. 
The ectopterygoid is toothless in the Pyrrhulinina but has conical 
teeth in the Lebiasini, and the bone is always proportionately larger 
than in the Nannostomina. The fenestra between the quadrate, 
symplectic, and metapterygoid is absent. 

In the Erythrininae, the mandibular and palatine arches are 
very similar to those of the Characinae. The fenestra between the 
metapterygoid, quadrate, and symplectic is well developed, and the 
symplectic is not exceptionally deep at its posterior end. In Hoplias 
the ectopterygoid is a long, thin, tooth-bearing bone extending from 
the upper part of the quadrate across the inferior surface of the 
palatine. Anterior to its forward boundary is a small autogenous 
movable piece of tooth-bearing bone that is connected closely to the 
ectopterygoid, Sagemehl (1885, p. 95) called this bone the accessory 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 145 

palatine. For reasons discussed below, I tenatively prefer to call 
this bone the accessory ectopterygoid. Along the lateral edge of the 
ectopterygoid and accessory ectopterygoid is a row of large, conical 
teeth. Medial to this row on both bones is a narrow patch of small, 
conic teeth. In Erythrinus and Hoplerythrinus the ectopterygoid 
extends from the quadrate to over the ventral surface of the palatine 
(autopalatine). The ectopterygoid in both bears a band of conic 
teeth. In Hoplerythrinus, but not in Erythrinus, small conic teeth 
are present on the ventral surface of the mesopterygoid. In a speci- 
men of Hoplerythrinus unitaeniatus, 115 mm. in standard length, 
from British Guiana, CAS(IUM) 12331, the mesopterygoid teeth are 
well developed and cover much of that bone's lower surface; however, 
a specimen from Peru, SU 35044, which is 120 mm. in standard 
length, has only a few scattered teeth over the mesopterygoid. 

Hoedeman (1950b) described a new genus and species in the 
Erythrininae, Pseuderythrinus rosapinnis, from Dutch Guiana. His 
new genus seems distinguishable from Hoplerythrinus only by the 
number and distribution of teeth on the palatal arch. He recorded 
a narrow band of teeth on the palatines and no teeth on the "ptery- 
goid" for Pseuderythrinus, and he stated that Hoplerythrinus has a 
broad patch of teeth on the palatine and that there are teeth on the 
pterygoid. Hoedeman's pterygoid is the ectopterygoid of the present 
account. Examination of a cleared and alizarin-stained specimen and 
other, nonstained specimens of Hoplerythrinus does not confirm Hoede- 
man's views concerning the placement of teeth in this genus. I find 
that Hoplerythrinus has teeth on the ectopterygoid, no teeth on the 
palatine, and it may have a broad patch of teeth on the mesoptery- 
goid. The mesopterygoid teeth may be numerous or few and they 
are usually fewer in small specimens. It seems likely that the speci- 
men, 128 mm. in standard length, forming the basis of Hoedeman's 
new genus, has as yet failed to develop a dense aggregation of mesop- 
terygoid teeth, and it seems likely that this genus is a synonym of 
Hoplerythrinus. The color and other characters of Pseuderythrinus 
rosapinnis are very much like those of Hoplerythrinus unitaeniatus; 
further investigations probably would indicate that, at most, P. 
rosapinnis is a subspecies of H. unitaeniatus. The type of Pseudery- 
thrinus rosapinnis needs reexamination. 

Hoedeman (1950b, p. 85) recorded palatine teeth present in Hoplias, 
Hoplerythrinus, and Erythrinus and stated that there are no teeth 
on the "pterygoid" in Erythrinus. This disagrees with what I found 
in the specimens at hand. 

No specimens of these genera were found to have palatine teeth 
attached to the palatine. This fact is difficult to determine and re- 
quires considerable and careful dissection, especially in Erythrinus 



146 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

and Hoplerythrinus. In these two genera the tooth-bearing ecoptery- 
goid (= the pterygoid of Hoedeman) is firmly attached to, but not 
fused with, the palatine. The anterior part of the ectopterygoid lies 
ventral to the palatine in these two genera and can be mistaken 
easily for palatine dental plates or dermopalatines. The autogenous 
tooth-bearing bone below the palatine in Hoplias has been described 
above. 

Determination of the homologies of the tooth-bearing elements 
associated with the autopalatine of the Erythrininae cannot be stated 
precisely from the examination of adult stages only. The interpre- 
tation tentatively adopted here for the morphological situation in 
Hoplerythrinus is based on the following: The ectopterygoid is pres- 
ent and often tooth-bearing in other characids. It is presumed here 
that the ectopterygoid bone grows anteriorly under the lower surface 
of the palatines and that the dermal tooth-bearing bones under the 
autopalatine are not dermopalatine elements that have fused to the 
ectopterygoid but are simply anterior extensions of the ectopterygoid. 
The striations and growth pattern of these tooth-bearing bones indi- 
cate they are anterior extensions of the ectopterygoid. 

The accessory palatine of Sagemehl (1885, p. 95) that occurs in 
Hoplias may be interpreted in two ways, either as a dermopalatine 
or dental element that has not fused to the ectopterygoid or as an 
autogenous, anterior piece of the ectopterygoid. Starks (1926, p. 
161) maintains that the accessory palatine is homologous with the 
dermopalatine of Amia. This problem cannot be resolved with the 
information at hand. 

Probably the presence of teeth or dental plates associated with the 
palatine, ectopterygoid, and mesopterygoid is a conservative featm-e 
not retained in most other characids; in this respect the tooth pattern 
of the Erythrininae would be more primitive than that of the other 
subfamilies of living characids. If this is correct, the separation of 
the Erythrininae from the rest of the characids might be suspected 
then to have taken place when the characids had living representa- 
tives much more primitive than those extant. In this connection, 
the morphology and development of the first infraorbital and its 
relationship to the antorbital in the Erythrininae and possible signifi- 
cance of the infraorbital in the phylogeny of characids and Ostario- 
physi should be investigated. 

Hyoid and branchial arches (fig. 8). — The hyoid arch of Nan- 
nostomus and Poecilobrycon is not unlike that of the Characinae, and 
the relative positions of the bones of the arches are the same. There 
are two branchiostegal rays associated with the ceratohyal and one 
with the epihyal. Members of the tribe Pyrrhulinini also have three 
branchiostegal rays, while members of the tribe Lebiasinini have four 



SOUTH AMERICAN CHARACID FISHES — ^WEITZMAN 147 

branchiostegal rays, three associated with the ceratohyal and one with 
the epihyal. All members of the Erythrininae have five branchiostegal 
rays, fom* on the ceratohyal and one on the epihyal. 

The branchial arches of the Erythrininae and Lebiasininae are like 
those of the Characinae and differ mainly in general proportions cor- 
related with different head shape. 

Hoedeman (1950a, pp. 17, 19, 25) stated that pharyngeal teeth are 
absent in three species of his Nannostominae. All specimens of the 
Nannostomina, however, that I have examined have small, conic 
pharyngeal teeth much like those illustrated for Poecilobrycon harrisoni 
(fig. 8). The gill rakers of members of the Lebiasininae are simple; 
however, in the Erythrininae the gill rakers are rather complex. The 
bony cores of those of the upper limb are truncate with small conical 
"teeth" along their distal margins while those of the lower limb are 
serially graduated from truncate ones anteriorly to elongate sharp- 
pointed laminae of bone posteriorly. The dorsal edges of these 
laminae bear small conic "teeth" while the anterior truncated gill 
rakers have "teeth" along their distal borders. 

Weberian APPARATUS (fig. 9). — The Weberian apparatus of the 
Lebiasininae and Erythrininae is, in all major respects, like that of the 
genus Brycon and the Characinae, differing only in a few aspects of 
shape and in the proportions of some of the parts. The tripus of 
Poecilobrycon harrisoni is slightly different in shape from that of 
Brycon meeki but still retains the basic characid structure. Its 
dorsal proximal portion is placed slightly higher and more anterior on 
the body of the third vertebra. In some specimens of the Nan- 
nostomina examined there are small contingencies of bone across 
the joint between the neural pedicel and the neural complex. 

Pectoral girdle (fig. 10). — The pectoral girdle of Poecilobrycon 
harrisoni is much like that of the Characinae, having a large medial 
coracoid lamina. The main body of the cleithrum, like that of the 
Characinae, is a sickle-shaped lamella; however, the large foramen 
between the coracoid and the cleithrum, so typical of the Characinae, 
is absent or at best only a slight opening. The supracleithrum lacks 
a sensory canal in all members of the Nannostomina and the post- 
temporal is not forked, the lower spinous process found in the 
Characinae being absent. The relationships of the postcleithrum, 
mesocoracoid, and scapula are not markedly different from those of 
the Characinae. 

In the Pyrrhulinina, sensory canals are lacking in the post-tem- 
poral and supracleithrum, and the post-temporal is not forked. The 
rest of the pectoral girdle is typical of that of characids. Unlike the 
Nannostomina, the Pyrrhulinina have a large foramen between the 
coracoid and the cleithrum. In speicmens of Lebiasina and Pia- 



148 PROCEEDINGS OF THE NATIONAL MUSEUM ^'°^- "^ 

hucina, the foramen between the coracoid and cleithrum is well de- 
veloped. The rest of the pectoral skeleton in these two genera is 
like that of the Characinae. 

The pectoral girdle of the Erythrininae is unique among characids. 
The lower limb of the cleithrum is long and slender. The lateral 
inferior lamella is much reduced in extent. The median coracoid 
lamella is extremely reduced in size, being a very narrow crest or 
actually absent. In all other characids that I have examined, the 
coracoid extends anteriorly to, or almost to, the anterior edge of the 
cleithrum. In the Erythrininae, the coracoid extends anteriorly for 
only about one-half the length of the lower limb of the cleithi'um, and 
the foramen between the cleithrum and coracoid is absent. The 
cleithrum and post-temporal of the Erythrininae contain segments of 
the laterosensory canals. 

Classification of the Characid Subfamilies 
Lebiasininae and Erythrininae 

The following classification is based primarily on the evidence 
presented in the foregoing section. 

SUBFAMILY LEBIASININAE EIGENMANN, 1910 

Definition: (1) Large fenestra between quadrate and metaptery- 
goid absent. (2) Two diverging lamellae on ventral surface of 
ethmoid very poorly developed. (3) Lateral posterior edge of frontal 
not articulated with spinous process of sphenotic. (4) Six infraorbital 
bones present; antorbital and first infraorbital separate ossifications. 
(5) Accessory ectopterygoid (or dermopalatine?) absent. (6) Lower 
limb of cleithrum moderately stout, not long and slender, its lateral 
inferior lamella large. (7) Coracoid lamella a large flat plate, reach- 
ing to near anterior border of cleithrum. (8) Anastomosis between 
supraorbital and dermopterotic canal present. (9) Parietal and 
epiphyseal branches of supraorbital canal reduced in length, parietal 
branch never reaching parietal bone. (10) Perforated lateral line 
scales, when present, confined to region anterior to dorsal fin. (11) 
Frontal fontanel always absent in adults. (12) Teeth well developed 
in both jaws, unicuspid or multicuspid. (13) Premaxillary with one 
tooth row. (14) Dentary usually with two rows of teeth. (15) Gape 
short, not reaching orbit, or, at most, only reaching posteriorly little 
beyond verticle with anterior margin of orbit. (16) Branchiostegal 
rays three or four. (17) Gill membranes partially united but free 
from isthmus. (18) Body elongate, often fusiform or with fairly 
blunt head; body rounded in cross section in region of dorsal fin and 
head, compressed in region of caudal peduncle. (19) Adipose fin 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 149 

present or absent. (20) Anal fin short-based, with 8 to 14 rays. 
Males often with specialized anal fin rays; these expanded in anterior- 
posterior plane and often thickened laterally. (21) Dorsal fin always 
in advance of anal fin, placed over pelvic fins. (22) Caudal fin 
rounded or forked. (23) Scales large, 6 to 7 horizontal rows between 
dorsal and pelvic fines, 18 to 30 scales in longitudinal series. 
The tribes of subfamily Lebiasininae are: 

Lebiasinini Eigenmann, 1910 
Pyrrhulinini Eigenmann, 1910 

TRIBE lEBIASININI EIGENMANN, 1910 

Noraenclatural note: Eigenmann (1910) was the first to base a 
family group name, Lebiasininae, on the genus Lebiasina. Hoedeman 
(1950b) was the first to propose the use of this family group name as 
a tribe. 

Definition: (1) Four branchiostegal rays. (2) Premaxillary divided 
into well-developed upper and lower rami. (3) Premaxillary teeth 
tricuspid, in one row. (4) Dentary with teeth in two rows, outer row 
teeth tricuspid, inner with simple, conic teeth. (5) Dentary without 
inferior deep notch or groove, ventral surface smooth. (6) Parietal 
branch of supraorbital sensory canal moderately well developed, 
reaching almost to parietal bone. (7) Supratemporal laterosensory 
canals present. (8) Dermopterotic laterosensory canals present. 
(9) Extrascapular laterosensory canals and bones present. (10) 
Supracleithrum with laterosensory canal. (11) Frontalpterotic joint 
present on surface of skull. 

The genera of tribe Lebiasinini are: 

Lebiasina Valenciennes (1846), with two species. 
Piabucina Valenciennes (1849), with six or seven species. 

Remarks: Lebiasina has been separated from Piabucina by the 
absence of an adipose fin in the former and its presence in the latter. 
This is apparently a poor character in these fishes, for, as Eigenmann 
(1923, p. 123) noted, some specimens of Lebiasina bimaculata and 
Lebiasina multimaculata occasionally have an adipose fin. In the 
specimens I have been able to examine, the anterior wall of the 
posterior division of the swimbladder in Lebiasina bimaculata and 
Piabucina festae is "cellular" while in Piabucina erythrinoides and Pia- 
bucina panamensis it is not. The tribe Lebiasinini needs revision at 
the specific and generic level. 

TRIBE PYRRHULININI EIGENMANN, 1910 

Nomenclatural note: Eigenmann (1910) was the first to base a 
family group name, Pyrrhulininae, on the genus Pyrrlndina. Hoede- 
man (1954a) was the first to propose the use of this family group 
name as a tribe Pyrrhulinini. 



150 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- "^ 

Definition: (1) Three branchiostegal rays. (2) Premaxillary only 
weakly, or not at all, divided into upper and lower processes. (3) 
Premaxillary teeth simple conic to hexacuspid, in one or two rows. 
(4) Dentary with teeth in two rows, uni- to hexacuspid. (5) Dentary 
with inferior notch below or deep groove continuous with foramen for 
mental ramus of mandibular branch of trigeminal nerve. (6) Parietal 
and epiphyseal branches of supraorbital laterosensory canal extremely 
short, usually not over one or, rarely, two millimeters long even in 
largest specimens of largest species; parietal canal not reaching parietal 
bone. (7) Supratemporal laterosensory canal absent. (8) Dermo- 
pterotic canal absent. (9) Extrascapular laterosensory canals and 
bones absent. (10) Supracleithrum without enclosed laterosensory 
canal. (11) Frontal-pterotic joint absent, pterotic being excluded 
from contact with frontal by sphenotic. 

The subtribes of tribe Pyrrhulinini are: 

Pyrrhulinina 
Nannostomina 

SUBTRIBE PYRRHULININA EIGENMANN, 1910 

Definition: (1) Premaxillary-ethmoid articulation a very slightly 
movable, syndesmotic (fibrous) joint. (2) In adults, posterior shaft 
of prevomer reaches parasphenoid. (3) Preopercle extends anteriorly 
to near articular. (4) Maxillary bone extends posteriorly to region 
of second orbital. (5) Premaxillary with unicuspid teeth. (6) Teeth 
of dentary unicuspid, in two rows. (7) Dentary not divided into two 
well-defined regions by a deep ventral groove continuous with mental 
foramen. Dentary with notch below region of mental foramen. 
(8) Inferior portion of post-temporal fossa extremely large, its antero- 
lateral border extending anterior to, or beyond, synchondral portion 
of joint between sphenotic and pterotic bones, anteriorly beyond 
vertical from anterior edge of auditory foramen. 

The genera of subtribe Pyrrhulinina are: 

Pyrrhulina Valenciennes (1846), with perhaps a dozen species 
Copeina Fowler (1906), with two or three species 
Copella Myers (1956a), with four or five species 

SUBTRIBE NANNOSTOMINA EIGENMANN, 1909 

Nomenclatural note: Eigenmann (1909a) was the first to base a 
family group name, Nannostomatinae, on the genus Nannostomus. 
The proper spelUng of this family group name should have been 
Nannostominae (Miller, 1897, p. 132). 

Definition: (1) Premaxillary-ethmoid articulation a loose movable 
syndesmotic (fibrous) joint connected by fairly long, interosseous 
ligaments, but premaxillary not protractile, synovial joint absent. 



SOUTH AMERICAN CHARACID FISHES — ^WEITZMAN 151 

(2) Posterior shaft of prevomer reduced or absent in adults and young, 
never reaching parasphenoid. (3) Preopercle not reaching near articu- 
lar, extending anteriorly only to lower posterior process of quadrate. 

(4) Maxillary confined to anterior region of first infraorbital bone. 

(5) Premaxillary teeth 1- to 6-cusped, usually 3- to 6-cusped, multi- 
cuspid teeth flattened and incisor-like. (6) Dentary with outer 
row of flattened, 3- to 6-cusped incisor-like teeth and one inner 
row of unicuspid conic teeth. (7) Dentary divided into two distinct 
regions by a deep inferior groove continuous with the mental foramen. 
(8) Lower portion of post-temporal fossa not enlarged, its anterolateral 
border well posterior to vertical from synchondral joint between 
sphenotic and pterotic, anterolateral border of post-temporal fossa 
behind vertical from anterior edge of auditory foramen. 

The genera of sub tribe nannostomina are: 

Nannostomus Glinther (1872), with five or six valid species 
Poecilobrycon Eigenmann (1909b), with three valid species 

Note that Nannobrycon of Hoedeman (1950a) is here considered a 
subgenus of Poecilobrycon and Archicheir Eigenmann (1909b) is con- 
sidered a synonym of Poecilobrycon. Archicheir minutus Eigenmann 
(1909b), the type of Archicheir, is based on the young of Poecilobrycon 
harrisoni Eigenmann (1909b). A review of the genera and species 
of the Nannostomina is in preparation. 

SUBFAMILY ERYTHRININAE GILL, 1858 

Definition: (1) Large fenestra between quadrate and metapterygoid 
present. (2) Two diverging lamellae on ventral surface of ethmoid 
weU developed. (3) Lateral posterior edge of frontal articulated 
with spinous process of sphenotic. (4) Six infraorbital bones present; 
first infraorbital apparently fused with antorbital. (5) Accessory 
ectopterygoid (or dermopalatine) present, ventral to autopalatine 
either as an autogenous element or fused with ectopterygoid. (6) 
Lower limb of cleithrum long and slender, its lateral inferior lamella 
small. (7) Coracoid lamella small or absent, not large, flat plate 
as in other characids, not reaching to near anterior border of cleithrum. 
(8) Anastomosis between supraorbital and dermopterotic latero- 
seusory canal absent. (9) Parietal and epiphyseal branches of 
supraorbital canals well developed, parietal canal reaching parietal 
bone. (10) Perforated lateral fine scales present, continued unin- 
terrupted to caudal fin. (11) Frontal fontanel always absent in 
adults. (12) Teeth well developed, unicuspid, conic in both jaws. 
(13) Premaxillary with one tooth row. (14) Dentary with anterior 
tooth row bordering length of jaw, inner tooth row confined to pos- 
terior portion of mandibular ramus. (15) Gape long, reaching 



152 PROCEEDINGS OF THE NATIONAL MUSEUM ^"^- "^ 

posteriorly beyond vertical from anterior border of orbit. (16) 
Branchiostegal rays five. (17) Gill membranes partially united but 
free from isthmus. (18) Body elongate, with blunt head and snout; 
body rounded in cross section in region of dorsal fin and head, com- 
pressed in region of caudal peduncle, (19) Adipose fin absent. 
(20) Anal fin short-based, with about 10 to 12 rays. (21) Dorsal fin 
always in advance of anal fin, usually over pelvic fins. (22) Caudal 
fin rounded in profile. (23) Scales moderate to fairly large, about 
30 to 45 in a lateral series, about 7 to 12 in horizontal rows between 
dorsal and pelvic fins. 
The genera of subfamily Erythrininae are : 

Hoplias Gill (1903), with one or two species 

Erythrinus Scopoli (1771), with one species 

Hoplerythrimis Gill (1895), with one species 

Pseuderythrinus Hoedeman (1950b), with one species, status uncertain 

Remarks: Of the characters listed above, 3-8 seem to be diagnostic 
for the subfamily, distinguishing its members from all other characids. 

Relationships of the Subtribe Naimostomina and its Close 
Relatives, the Pyrrhulinina 

The classification above clearly indicates the close relationship of 
the Nannostomina and Pyrrhulinina. The unique structure of the 
dentary in these two groups, together with the accumulation of such 
characters as three branchiostegal rays, a premaxillary without a clear 
upper or lower process, short parietal and epiphyseal laterosensory 
canals, no supratemporal or dermopterotic canals, no extrascapular 
bone or canal, no canal embedded in the supracleithrum, and the lack 
of a frontal-pterotic joint, cannot be due to convergent evolution 
but only to a common ancestry. Further inspection of these fishes 
reveals other similarities; for example, their scales are almost identical 
in shape and structure, both groups have members with modified anal 
fin rays in the males, both have a tendency to lose canals in orbital 
bones, and one species, Nannostomus espei, has a color pattern re- 
markably like that of Pyrrhulina vittata and Pynhulina spilota. 
Nannostomus espei also has scales that more closely approach the 
shape of those of Pyrrhulina than the scales of any other species in 
the Nannostomina. The osteology of Nannostomus espei, however, 
is typically that of Nannostomus. 

The Nannostomina are specialized for feeding on small animal 
organisms that occur on plants, rocks, and other objects. They also 
feed on slowly moving free-living organisms. Their movements are 
relatively slow and precise. The Pyrrhulinina are more active and 
more predaceous, having larger mouths, attacking larger prey, and 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 153 

striking their prey harder. The jaws of the Nannostomina are far 
more speciahzed than those of the Pyrrhulinina, The large post- 
temporal fossa in the Pyrrhulinina is an interesting specialization. 
The large amount of muscle tissue found in this area undoubtedly 
is associated functionally with feeding and swimming behavior, but 
this needs further study. 

In addition to the differences between the Pyrrhulinina and 
Nannostomina noted in the classification above, these two groups 
have a consistent difference in body shape. The body form in the 
Nannostomina usually is smoothly fusiform, whereas that in the 
Pyrrhulinina is not, the profile showing a slight notch behind the 
dorsal fin, the belly being more or less flat, and the back being less 
arched. Of all the species in the Nannostomina, Nannostomus espei 
has a body shape closest to that of the Pyrrhulinina. 

Despite the differences between the Pyrrhuhnina and the 
Nannostomina, I believe they may be united justifiably in the tribe 
Pyrrhulinini, based on the morphological characters listed above. 

The relationships of the Pyrrhulinini are, without doubt, with 
the Lebiasinini, and these two tribes form the subfamily Lebiasininae, 
The characters Usted in the classification for this subfamily (p. 148) 
are held in combination by no other characids. A full discussion of 
the relationships between the Lebiasinini and the Pyrrhulinini should 
await fuller treatment of the genera in these two tribes. 

From the information at hand it seems safe to conclude that the 
Pyrrhulinini were derived from characids somewhat similar perhaps 
to the existing members of the Lebiasinini and that the Pyrrhulinina 
and the Nannostomina had a common ancestor. Neither of these 
two subtribes can be derived from the other, but their common 
ancestor must have differed from the recent members of the Lebiasinini 
in the structure of the jaws and reduction of the laterosensory system 
of the head. 

Relationships of the Lebiasininae and Erythrininae 

What are the relationships of the Lebiasininae and Erythrininae 
that can be determined from the present osteological study? Con- 
sidering the absolute lack of fossil evidence in this case, inferences 
about phylogenetic relationships based on recent material are bound 
to bear only a vague resemblance to the actual course of evolution; 
nevertheless, an attempt should be made to indicate relationships 
even if this involves nothing more than noting that certain groups 
appear associated because they have a certain number of characters 
in common. 



154 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- ^^^ 

The relationships of the Erythrininae to other recent characids 
remain rather obscure. They are not close to the Characinae and 
they do not appear derived from them. A consideration of the primi- 
tive versus specialized aspects of their skeleton may give some hint 
regarding their relationships. In general, a reduction in number of 
branchiostegal rays in teleosts can be considered a specialized feature 
(Hubbs, 1919). In all probability, the presence of five branchiostegal 
rays in the Erythrininae is relatively primitive for characids. The 
absence of the supraorbital in both the Erythrininae and the Lebiasin- 
inae probably is specialized and I suspect that this is also true for 
the morphology of the antorbital in the Erythrininae. Dental 
plates and teeth associated with the palatine, ectopterygoid, and 
mesopterygoid in the Erythrininae is probably a primitive feature 
in these fishes. The unique absence of a connection between the 
dermopterotic and supraorbital laterosensory canals in the Eryth- 
rininae probably is specialized. Whether the articulation between 
the sphenotic and frontal in the Erythrininae is primitive or specialized 
cannot be said. It would seem that the pectoral girdle of the Eryth- 
rininae is rather specialized because the interosseus space, usually 
present in teleosts and other subfamihes of characids (Starks, 1930, 
p. 90), is absent; however, the gross morphology of the pectoral 
girdle of the Erythi'ininae is in some respects similar to that of Amia 
and it is possible that the form of the pectoral girdle in the Erythrin- 
inae is rather primitive. Likewise, it is difficult to evaluate the 
generalized (versus specialized) nature of other characters of the 
Erythrininae. The short-based anal fin is probably primitive with 
regard to the Characidae and Cypriniformes as a whole, and the same 
may be true of the large scales. It is also possible that the rather 
blunt, cylindi'ical body shape is primitive. 

Examination of the osteology of Hepseius indicates certain relation- 
ships with Erythrinus and Hoplias. Bertmar (1959, p. 350; and in 
litt.) has noted that Hepsetus is more primitive from the embryo- 
logical point of view than the other ten characids he examined and 
that all the characids he investigated are more primitive in certain 
respects than Amia. This of course does not mean that characids 
are more primitive than Amia, but it does pose questions about the 
origins of the Cypriniformes. The Erythrininae appear to be more 
primitive osteologicaUy than Hepsetus. Investigation of the ontogeny 
of their chondrocranium should be very fruitful. 

About aU that can be concluded here is: (1) the Erythrininae are 
definitely characid in morphology; (2) but their basic structure is 
more remote from that of the so-called "central gi'oup of characids" 
(i.e., the Characinae) than of any other known living characid; (3) 
they probably were not derived from a characid ancestor that was 



SOUTH AMERICAN CHARACID FISHES — ^WEITZMAN 155 

like any living member or members of the Characinae. There is per- 
haps more reason to give separate family status to the Erythrininae 
than to any other subfamily of characids. Also there is some reason 
to suspect that at least in certain featm-es, the Erythrininae are the 
most archaic of living characids. In other aspects, however, they 
are possibly quite specialized. 

What are the relationships of the subfamily Lebiasininae? Super- 
ficially, members of the tribe Lebiasinini look much like members of 
the Erythrininae because of their large scales, short-based anal fin, and 
the overall similar appearance of the body and head. In addition, both 
groups lack the supraorbital. The last is possibly not too important 
a character, for loss of an element probably never should be consid- 
ered as important as the development of a new structure. Except 
for the fact, however, that these two groups are characids and have 
similarly shaped cranial bones because of their similar general body 
and head shape, they have little in common osteologically. In aU the 
unique respects in which the Erythrininae differ from the Characinae, 
the Lebiasininae are like the Characinae. As noted above, members 
of the tribe Lebiasinini and indeed the entire Labiasininae have a 
few very unique characters that are different from those of both the 
Characinae and the Erythrininae. The unique lack of a fenestra 
between the quadrate and metapterygoid appears specialized. Also, 
the general reduction in the laterosensory canals of the head of the 
Lebiasininae and the unique laterosensory head canals of the member 
subtribes of the Pyrrhulinini are very specialized. It seems likely 
(1) that the Lebiasininae are somewhat more closely related to the 
Characinae than to the Erythrininae and (2) that the Lebiasininae 
evolved from a common characid stock that gave rise to Lebiasininae 
and Characinae but not to the Erythrininae. 

According to several authors, one of the distinguishing characters 
of the Lebiasinini is the possession of a "cellular" anterior wall of the 
posterior chamber of the swimbladder. The present investigation 
indicates that certain species (Piabucina panamensis and P. ery- 
thrinoides) in this group lack this feature while others (Lebiasinina 
bimaculata and Piabucina jestae) do have it. Obviously, the group 
needs revision and, since Bohlke (1958, p. 94) has indicated he is doing 
this, no revision is attempted here. It should also be noted that, of 
the Erythrininae, at least Hoplerythrinus unitaeniatus and Erythrinus 
erythrinus have a cellular anterior wall of the posterior chamber of 
the swimbladder. Hoplias malabaricus does not. The significance 
of the similar "cellular" structure of swimbladders of some members 
of the Erythrininae to that of some of the Lebiasinini is difficult to 
assess. In view of the distinct osteological differences between the 
Erythrininae and the Lebiasinini, however, I prefer to believe that 



156 PROCEEDINGS OF THE NATIONAL MUSEUM ''^^- "^ 

this equivalence of structure is due to similar adaptation to stagnant 
water and that it does not indicate close genetic relationships. Os- 
teological comparisons indicate that the Lebiasininae are not related 
closely to such genera as Anostomus, Leparinus, and Hemiodus. 

Summary 

The primary results of this study are as foUows: The Lebiasininae 
and Erythrininae are defined and their relationships to each other 
and to other characids are discussed. The Lebiasininae are not 
related closely to the Erythrininae as assumed by many earlier authors. 
The Erythrininae cannot have been derived directly from ancestors 
that were like recent members of the Characinae (as defined by 
Weitzman, 1962, p. 48). Although the evidence is inconclusive, of 
all living characids, the Erythrininae are probably the most conserva- 
tive (in some ways). It is sm^mised that the Characinae and Erythri- 
ninae were derived from a common characid stock remote in time and 
morphology from these two subfamiHes as they are known today. 
The Lebiasininae, in many important respects, are more closely 
allied morphologically to the Characinae than to the Erythrininae 
despite their superficial resemblance to the Erythrininae. It appears 
that the Lebiasininae may have been derived from early members 
of the Characinae or at least from ancestral characids that were more 
like the Characinae than the Erythrininae in their morphology. 

The subfamily Lebiasininae consists of two tribes, the Lebiasinini 
and the Pyrrhulinini. Both are defined herein. The Lebiasinini 
needs revision, but no attempt was made to reevaluate in this study 
its genera and species. The Pyrrhulini consists of two subtribes, 
the Pyrrhulinina and the Nannostomina, and both are defined herein. 



Literature Cited 

Bertmar, Gtjnnar 

1959. On the ontogeny of the chondral skull in Characidae, with a discussion 
on the chondrocranial base and the visceral chondrocranium in 
fishes. Act. Zool. Stockholm, vol. 40, pp. 203-364. 
BoHLKE, James 

1958. Studies of fishes of the family Characidae. — No. 14. A report on 
several extensive recent collections from Ecuador. Proc. Acad. 
Nat. Sci. Philadelphia, vol. 110, pp. 1-121, pis. 1-7. 

BOULENGER, GeoRGE 

1904. Fishes (systematic account of Teleostei) . /n S. F. Harmer and A. E. 
Shipley, The Cambridge Natural History, vol. 7 (xviii4-760 pp.), 
pp. 539-727. 
Branson, Branlet A., and Moore, George A. 

1962. The lateralis components of the acoustico-lateralis system in the sun- 
fish family, Centrarchidae. Copeia, no. 1, pp. 1-108. 



SOUTH AMERICAN CHARACID FISHES — WEITZMAN 157 

COCKERELL, ThEODORE D. A. 

1914. The scales of the South American characinid fishes. Ann. Carnegie 

Mus., vol. 9, pp. 92-113, pis. 23-28. 
Cuvier, Georges L. C. F. D., and Valenciennes, Achille 
1846. Histoire naturelle des poissons, vol. 19, xix-f549 pp. 
1849. Histoire naturelle des poissons, vol. 22, xx+92 + 332 pp. 
Eigenmann, Carl H. 

1909a. The fresh-water fishes of Patagonia and an examination of the Archi- 

plata-Archhelenis theory. Rep. Princeton Univ. Exped. Patagonia, 

1896-1899, vol. 3, Zool., pp. 225-374, pis. 30-37. 
1909b. Some new genera and species of fishes from British Guiana. In 

Reports on the expedition to British Guiana of the Indiana Univer- 
sity and the Carnegie Museum, 1908, report no. 1. Ann. Carnegie 

Mus., vol. 6, pp. 4-54. 
1910. Catalogue of the fresh- water fishes of tropical and south temperate 

America. Rep. Princeton Univ. Exped. Patagonia, 1896-1899, 

vol. 3, Zool., pp. 375-511. 
1912. The fresh-water fishes of British Guiana. Mem, Carnegie Mus., vol. 

5, xxii + 578 pp., 103 pis. 
1923. The fishes of Western South America, pt. 1. Mem. Carnegie Mus., 

vol. 9, pp. 1-346, pis. 1-35. 
Eigenmann, Carl H., and Eigenmann, Rosa Smith 

1889. A review of the Erythrininae. Proc. California Acad. Sci., ser. 2, 

vol. 2, pp. 100-116, 1 pi. 
1891. A catalogue of the fresh-water fishes of South America. Proc. U.S. 

Nat. Mus., vol. 14, pp. 1-81. 
Fernandez- Yepez, Augustin 

1947. Charaxodon, a new genus of characid fishes from South America. 

Evencias, Estab. Venezolano Cien. Nat., ser. 1, no. 3, 3 pp. 
1955. Los peces neotropicales de la familia Acestrorhynchidae. Rev. Fac. 

Agric. Univ. Central Venezuela, vol. 1, pp. 449-459. 
Fowler, Henry W. 

1906. Further knowledge of some heterognathous fishes, part 1. Proc. 

Acad. Nat. Sci. Philadelphia, vol. 58, pp. 293-352. 
1950. Os peixes de dgua doce do Brasil (2.» entrega). Arq. Zool. Est. Sao 

Paulo, vol. 6, pp. 205-404. 
Gill, Theodore Nicholas 

1958. Synopsis of the fresh-water fishes of the western portion of the Island 

of Trinidad, W. I. Ann. Lye. Nat. Hist. New York, vol. 6, pp. 

363-430. 
1895. The differential characters of the characinoid and erythrinoid fishes. 

Proc. U.S. Nat. Mus., vol. 18, pp. 205-209. 
1903. Note on the fish genera named Macrodon. Proc. U.S. Nat. Mus., 

vol. 26, pp. 1015-1016. 
Gregory, William King, and Conrad, George Miles 

1938. The phylogeny of the characin fishes. Zoologica (New York), vol. 

23, pp. 319-360. 
GtJNTHER, Albert C. L. G. 

1864. Catalogue of the fishes in the British Museum, London, vol. 5, xxii-f 

455 pp. 
1872. On a new genus of characinoid fishes from Demerara. Proc. Zool. 

Soc. London, 1872, p. 146. 



158 PROCEEDINGS OF THE NATIONAL MUSEUM ^^^- "^ 

HOEDEMAN, J. J. 

1950a. Rediagnosis of the characid-nannostomine fish genera Nannostomus 
and Poecilobrycon. Amsterdam Natur., vol. 1, pp. 11-27, pis. 8-9. 

1950b. A new characid-Erythrinine fish {Pseuderythrinus rosapinnis gen. et 
sp. nov.). Amsterdam Natur., vol. 1, pp. 79-91. 

1954a. Notes on the ichthyology of Surinam (Dutch Guiana), 3: A new- 
species and two new subspecies of Nannostoraidi from the Surinam 
River. Beaufortia, Ser. Misc., Publ. Zool. Mus., vol. 4, pp. 81-89. 

1954b. Aquariumvissen-encyclopaedie, nieuwe importen, 527 pp. 

1956a. fitude des ecailles des poissons de I'ordre des Cypriniformes (pt. 
1 — particulierement les characoides) . L' Aquarium et les Poissons, 
6"* anee, pp. 9-19. 

1956b. Aquariumvissen-encyclopaedie, nieuwe importen, pp. 528-728 [sup- 
plement to 1954b.] 
HuBBS, Carl L. 

1919. A comparative study of the bones forming the opercular series of 
fishes. Journ. Morph., vol. 33, pp. 61-71. 
Meinken, Hermann 

1931. Ueber die Entwicklung der Flossen in den Jugend stadien ver- 
schiedener Fische. Blatt. Aquar.-Terrarienkunde, Jahrg. 42, vol. 
17, 18, 19, pp. 1-14. 
Miller, Walter 

1897. Scientific names of Latin and Greek derivation. Proc. California 
Acad. Sci., ser. 3, Zool., vol. 1, pp. 115-143. 
Myers, George S. 

1956a. Copella, a new genus of pyrrhulinin characid fishes from the Amazon. 
Stanford Ichth. Bull., vol. 7, pp. 12-13. 

1956b. Esomus rehi, an Indo-Malayan cyprinid fish. Stanford Ichth. Bull., 
vol. 7, pp. 13-14. 
PrroN, L. 

1938. Les Characinidae fossiles de Menat (P.-de-D.) . Rev. Sci. Bourb. Cent. 
France, Moulins Aout-Decembre 1938, nos. 3-4, pp. 98-104. 
Regan, Charles Tate 

1906. On the fresh-water fishes of the Island of Trinidad, based on a col- 
lection, notes, and sketches made by Mr. Lechmere Guppy. Journ. 
Proc. Zool. Soc. London, 1906, pp. 378-393. 

1911. The classification of the teleostean fishes of the order Ostariophysi, 

1 : Cyprinoidea. Ann. Mag. Nat. Hist., ser. 8, vol. 8, pp. 13-32, pi. 2. 

1912. A revision of the South American characid fishes of the genera Chal- 

ceus, Pyrrhulina, Copeina and Pogonocharax. Ann. Mag. Nat. 
Hist., ser. 8, vol. 10, pp. 387-395. 
Sagemehl, M. 

1885. Beitrage zur vergleichenden Anatomie der Fische, III: Das Cranium 
der Characiniden nebst allgemeinen Bemerkungen iiber die mit 
einem Weber'schen Apparat versehenen Physostomenfamilien. 
Morph. Jahrb., Bd. 10, pp. 1-119, pis. 1-2. 
ScopoLi, Giovanni Antonio 

1777. Introductio ad historiam naturalem . . . Prague, x -1-506 + 34 pp. 
Starks, Edwin 

1926. Bones of the ethmoid region of the fish skull. Stanford Univ. Publ., 

Univ. Ser. Biol. Sci., vol. 4, pp. 139-338. 
1930. The primary shoulder girdle of the bony fishes. Stanford Univ. 
Publ., Univ. Ser. Biol. Sci., vol. 6, pp. 149-239. 



SOUTH AMERICAN CHARACID FISHES WEITZMAN 159 

Steindachner, Franz 

1876. Ichthyologische Beitrage (V), II: Ueber eiiiige neue Fischarten, 

insbesondere Characinen und Siluroiden aus dem Araazonenstrome, 

Sitzb. Akad. Wiss. Wien, math.-nat. Classe, vol. 74, pp. 49-240, 

15 pis. 
Valenciennes, Achille — see Cuvier, Georges, and Valenciennes, Achille. 
Weitzman, Stanley H. 

1954. The osteology and relationships of the South American characid 

fishes of the subfamily Gasteropelecinae. Stanford Ichth. Bull., 

vol. 4, no. 4, pp. 212-263. 
1960. The systematic position of Piton's presumed characid fishes from the 

Eocene of central France. Stanford Ichth. Bull., vol. 7, no. 4, 

pp. 114-123. 
1962. The osteology of Brycon meeki, a generalized characid fish, with an 

osteological definition of the family. Stanford Ichth. Bull., vol. 8, 

no. 1, pp. 3-77. 



728-^26—64- 



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SOUTH AMERICAN CHARACID FISHES — WEITZMAN 169 

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U.S. GOVERNMENT PRINTING 0FFICEtl964 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1964 Number 3500 



THREE NEW SPECIES OF FROGFISHES 

FROM THE INDIAN AND PACIFIC OCEANS 

WITH NOTES ON OTHER SPECIES 

(FAMILY ANTENNARIIDAE) 



By Leonard P. Schultz 



The purposes of this paper are: (1) to evaluate further certain 
diagnostic characters of frogfishes; (2) to call attention to nomencla- 
tural changes; (3) to describe three new species; (4) to record speci- 
mens and data not available at the time I reviewed the family 
Antennariidae (Proc. U.S. Nat. Mus., vol. 107, no. 3383, pp. 47-105, 
8 figs., 14 pis., 1957). 

I am grateful to the staffs of the California Academy of Sciences 
(CAS), San Francisco, California; of the University of Hawaii (UH), 
Honolulu, Hawaii; and of the Division of Systematic Biology, Stanford 
University (SU), for permitting me to study the specimens in their 
collections. The following abbreviations, refer to other specimens 
recorded in this paper: RMNH (Rijikmuseum van Natuurlijke 
Historic, Leiden) and USNM (United States National Museum). 

171 

729-369—64 



172 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

Diagnostic Characters 

The study of additional material, not available previously, has 
enabled me to reconsider certain unsolved problems as well as to in- 
vestigate new ones. My description of the bait as a ''non-filamentous 
simple slender tentacle" has led to misinterpretation. This type of 
bait is considered to be a simple or unbranched central stalk, varying 
from a single slender threadlike tentacle to one that is somewhat 
fleshy, as contrasted to a tuft of tentacles or a bulbous base with ten- 
tacles. In addition, it should be understood that the bait is extensible 
and, when contracted, may be fleshy in nature; yet it retains the single 
central stalk. 

Further attention has been given to the evaluation of the presence 
or absence of warts on the skin of frogfishes as a taxonomic character. 
Two species with warts were described by Bleeker, Antennarius 
phymatodes and A. oligospilos. During my examination of specimens 
of A. moluccensis Bleeker in the collection of the University of Hawaii, 
I encountered an abnormal specimen with "warts" — 'UH 403, collected 
at Diamond Head, Oahu, measuring 87 mm. from tip of snout to base 
of caudal fin. Dr. Victor G. Springer informed me that he investi- 
gated "warts" on a large frogfish collected in the Gulf of Mexico and 
that the warts were encysted nematodes. Thus, "warts" on the skin 
of frogfishes may be of questionable value in distinguishing species. 

Briggs (Copeia, vol. 2, p. 440, 1962) distinguished Antennatus reticu- 
laris from A. strigatus on supposed differences in the length of the 
ilicium in relation to the length of the second dorsal spine and on the 
characteristic that the esca is blunt and lobed instead of lanceolate. 

Rosenblatt (Copeia, vol. 2, pp. 462-464, 1963) investigated this 
problem by measuring the length of the ilicium and the second dorsal 
spine on 62 specimens, then analyzing his data statistically. He con- 
cluded that "only one population was represented in the sample." 
He compared the pectoral ray counts of A. strigatus and A. bigibbus 
and concluded that the eastern Pacific population of A. strigatus was 
distinct from the western Pacific population of A. bigibbus. This 
leaves the two species as valid and in agreement with the conclusions 
of Schultz (1957). 

The importance of fin ray counts in the identification of frogfishes 
is evident when one studies table 1. I have added, therefore, the 
new counts made on the specimens listed below to my table 1 published 
in 1957. Additional counts are recorded from other specimens men- 
tioned elsewhere in this paper as well as some furnished by Dr. Richard 
Rosenblatt. 

Table 1 includes the nomenclatural changes discussed in this report. 



NEW INDO-PACIFIC FROGFISHES — SCHULTZ 



173 



Specimens Examined 

Antennarius aitipinnis 

SU 30045, Taruna, Sangi Island, Sangir Islands, Herre, June 24, 1928, 31 mm. 
SU 27238, Sitankai, Sulu, Philippines, Herre, August 7, 1931, 32 mm. 
SU 27239, Dumaguete, Philippines, Herre, 19 mm. 

Antennarius avalonis 

CAS [no number], Guaymas, Sonora, Mexico, February 1951, 200 mm. 

CAS 6627, latitude 0°55'S., longitude 90°31'W., Hancock Expedition, dredge 
68-60 fathoms, Jan. 26, 1934, 68 mm. 

CAS [no number], Guaymas, Mexico, D. A. Simpson, April 2, 1946, 3 speci- 
mens, 32-50 mm. 

SU 9979, Santa Catalina Island, California, holotype of A. avalonis Jordan 
and Starks, 267 mm. 

SU 16586, San Carlos Bay, Sonora, Mexico, June 1, 1950, 44 mm. 

SU 14968, Rocky Bluff Bay, Sonora, Mexico, 13 mm. 

Table 1. — Counts recorded for species of Antennariidae 



Genera, subgenera, and 
species 



Number of fin rays 



Soft dorsal 



11 12 13 14 15 16 



Anal 



6 7 



Pcctoralt 



7 8 



10 11 12 13 14 



Tathicarptis 

butlen* 

bulleri 
TrichophTyne 

rosace lis 

mitchelli* 
Nudiantennarius 

tubteres 
Abantennarius 

duescus 

analis 
Rhycherus 

filamentosus* 
Histiophryne 

bougainviUi* 

scortea* 
Echinophryne 

crassispina* 

glauerti* 
Phrynelox 
Phrynelox 

striatus* 

striatus 

melas* 

scaber 

lochites, new species 
Triantennatus 

cunninghami 

zebrinus 

atra 

nox 

tridena 



3 - - 

1 - - 



3 - 

1 - 



1 1 - 
- 2 1 



4 - 
1 2 



- 3 - - 



- 3 

- 3 



1 1 
- 1 



3 - 



1 1 



2 - 

- 3 

1 2 

1 2 

- 1 



- 1 

- 2 

- 2 
1 49 

- 4 



1 - 

1 1 

1 - 

50 1 

4 - 



2 - 
5 61 
2 6 



1 - 

6 - 
5 - 

7 - 
42 1 



1 1 
- 10 



43 



1 12 - 

7 66 4 



174 PROCEEDINGS OF THE NATIONAL MUSEUM vc 

Table 1. — Counts recorded for species of Antennariidae — Continued 



Genera, subgenera, and 
species 



Antennatus 
Antennatus 

bigibbus 

strigatus 
Golem 

cryptacanlhus 
LopMocharon 
Lophiocharon 

caudimaculatus 
Uniantennatus 

campylacanthus 

horriduf 

horridus* 

tenebTosus* 

tenebrosus 
Antennarius 
Fowlerichthys 

Tadiosus 

radiosus* 

avalonis 

sarasa* 

ocellatus 
Antennarius 

hispidus 

moluccensis* 

moluccensk 

leucosoma* 

pardalis 

chiTonede-s 

muUiocellatus 

phymatodes 

japonicus, new species 

indicus, new species 

sanguineus 

drombus 

cocdneus 

bermudensis 

dorehensis* 

notophihalmus* 

notophthalmus 

verrucosus 

altipinnis 

pauciradiatus 

nummifer 
Histrio 

histrio 



Number of fin rays 



Soft dorsal 



11 12 13 14 15 16 



1 29 



1 79 6 



29 1 

1 - 

17 - 



1 21 

- 24 

- 2 

- 1 

- 3 

- 4 

- 2 
1 20 

- 13 

- 22 



3 32 2 - 



Anal 



Pectoral t 



10 7 



9 10 11 12 13 14 



12 1 



- 28 2 - - 



1 - 
3 1 



30 1 
1 - 

17 - 



-59- 



- - ?8 
2 13 - 
1 6 - 

- 4 - 

- 1 - 



- 1 - 
-62 

- 24 1 
1 22 - 



19 3 - - 

12 1 - - 

22 - - - 

30 2 - - 



- - 2 27 - 

- 1 13 2 - 



46 2 



2 
2 1 



- - 107 5 

- 4 51 5 

- - 59 - 

- - 1 - 
3 26 3 



9 17 
1 - 



?1 - 

27 - 

13 - 

6 - 

2 - 



- - 13 3 

- - 12 36 
3 43 1 - 
2 2 - - 
1 - - - 



- 3 1 

- 7 1 



- 42 5 - - - - 
2 21 1 - - - - 

- - 34 4 - - - 

- 2 72 4 - - - 



*Counts recorded from literature. 

tCounts for both pectorals included when the number exceeds the count for 
the anal or dorsal ravs. 



NEW INDO-PACIFIC FROGFISHES — SCHULTZ 175 

Antennarius chironectes 

CAS [no number], Honolulu, E. M. Ehrhorn, 1915, 18 mm. 

CAS 6371, Honolulu, E. M. Ehrhorn, 92 mm. 

CAS 7400, Honolulu, A. Scale, 1935, 93 mm. 

SU 20463, Cagayancillo, Philippines, 63 mm. 

SU 8468, Honolulu, Albatross, 1902, 2 specimens, 52 and 101 mm. 

SU 8439, Laysan Island, holotype of A. laysanins Jordan and Snyder, 69 mm. 

UH 2338, Waiamae, Oahu, Aug. 12, 1955, 150 mm. 

Antennarius coccineus 

CAS [no number], Pago Pago, Samoa, A. Seale, May 1929, 72 mm. 
CAS [no number], Pago Pago, Samoa, A. Seale, May 1929, 70 mm. 
UH 1159, Hull Island, Phoenix Islands, July 12, 1950, 74 mm. 

Antennarius dromhus 

UH 1775, Hauula Park, Oahu, A. Tester, Jan. 21, 1953, one specimen 

UH 531, Waikiki, Oahu, Cuttress, April 4, 1950, one specimen 

UH 1832, Diamond Head, Oahu, one specimen 

UH 1227, Waialea School, Oahu, Feb. 4, 1951, two specimens 

UH 816, Diamond Head, Oahu, May 16, 1950, two specimens 

UH 1995, Waimea, Oahu, July 25, 1955, 62 mm. 

UH 338, Hauula Park, Oahu, June 28, 1949, 26 mm. 

UH 1751, Waikiki, Oahu, 40 mm. 

Antennarius multiocdlatus 
SU 52342, Port of Fortaleza, Brazil, Mucuripe, Feb. 23, 1945, 50 mm. 

Antennarius moluccensis 

CAS [no number], Hawaii, Thayer, Sept. 23, 1909, HI mm. 

CAS [no number], Hawaii, 173 mm, 

SU 8455, Honolulu, 53 mm. 

SU 8458, Honolulu, Albatross, 1902, 84 mm. 

UH 148, off Oahu, 162 mm. 

UH 1995, Waimea, Oahu, July 25, 1955, 36 mm. 

UH 268, Waimea, Oahu, Feb. 22, 1949, 1 specimen 

UH 403, Diamond Head, Oahu, Dec. 15, 1949, 87 mm. 

Antennarius nummifer 
SU 7600, Misaki, Japan, holotype of A. sanguifluus Jordan, 44 mm. 

Antennaries notophthalmus 

SU 27241, Unisan, Tayabas Province, Luzon, Philippines, Herre, Feb. 15, 
1924, 2 specimens, 52 mm. 

Antennarius sanguineus 

SU 14967, Cleopha Island, Tres Marias Islands, Mexico, Feb. 23, 1940, 
31.5 mm. 



176 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i] 

Antennaius bigibbus 

SU 27236, Dumaguete, Philippines, Herre, June 20-26, 1931, 2 specimens, 

23 and 34 mm. 
SU 29562, Mabini, Batangas, Philippines, Herre, March 1933, 34 mm. 
SU 8461, Honolulu, O. P. Jenkins, Albatross, 3 specimens, 15 and 19 mm. 
SU 3247, Honolulu, O. P. Jenkins, 5 specimens, 19 and 24 mm. 
UH 338, Hauula Park, Oahu, June 28, 1949, 33 mm. 
UH 2100, three miles west of Lahaina, Maui, Aug. 5, 1955, 33 mm. 

Aniennatus strigatus 

CAS W56-236, Clipperton Island, northeast side, Oct. 22-29, 1956, 34.3 mm. 
CAS W58-296, Clipperton Island, Aug. 20, 1958, 20 mm. [abnormal dorsal 

fin with last 4 rays branched] 
CAS W58-291, Clipperton Island, west end, Aug. 16, 1958, 22 mm. 

Lophiocharon caudimaculatus 

CAS [no number or locahty], 5 specimens, 57 to 92 mm. 
SU 39498, Singapore, Herre, Oct. 10, 1940, 2 specimens, 45 and 83 mm. 
SU 30651, Singapore, Herre, March 14, 1934, 4 specimens, 39 to 81 mm. 
SU 27872, Sandaken, British North Borneo, Herre, July 3, 1929, 3 specimens, 

65 to 87 mm. 
SU 32738, Singapore Harbor, Herre, May 7, 1937, 3 specimens, 53 to 69 mm. 
SU 35781, Singapore Market, Herre, May 1937, 2 specimens, 84 and 85 mm. 
SU 30652, Singapore, Herre, 2 specimens, 52 and 80 mm. 
SU 20204, Cuyo, Philippines, R. C. McGregor, holotype of A. lithinostomus 

Jordan and Richardson, 84 mm. 

Phrynelox atra 

CAS [no number or locality, identification uncertain], 82 mm. 
SU 9234, Port Jackson, Australia, 1 specimen 
SU 3259, Honolulu, Jenkins, 1889, 60 mm. 

Phrynelox nox 

SU 7601 Japan, cotype of A. nox Jordan, 61 mm. 

SU 7603, Nagasaki, Japan, holotype of A. no2 Jordan, 84 mm. 

SU 7599, Misaki, Japan, two cotypes of A. nox Jordan, 58 and 60 mm. 

Phrynelox scaber 
CAS 8819, Green Turtle Bay, Bahamas, 67 mm. 

Phrynelox striatus 

SU 9138, Port Jackson, Australia, 106 mm. 

SU 20725, Lake lUawarra, Australia, 87 mm. [bait is abnormal with 5 
branches] 



Proc us. Nat mus. Vol. 116 



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NEW INDO-PACIFIC FROGFISHES — SCHULTZ 177 

Phrynelox tridens 

CAS [no number], Tateyama Bay, Chiba Prefecture, Japan, Terazaki, Aug. 

29, 1946, 59 mm. 
CAS [lU 10680] Misaki, Sagami, Japan, Jordan and Snj'der, 3 specimens, 

49 to 52.5 mm. 
SU 32414, Tinghai, Chusan Island, Cheking Province, China, 2 specimens, 

47 and 63 mm. 
SU 25636, Nagasaki, Japan, 39 mm. 
SU 7227, Nagaski, Japan, 18 specimens, 27 to 59 mm. [one specimen has 

4 tentacles] 
SU 7228, Wakanoura, Japan, 7 specimens, 53 to 78 mm. 
SU 23424, Wakanoura, Japan, 5 specimens, 42 to 59 mm. 

Nomenclatural Changes 

Three new frogfish names have been pubhshed since my review. 
Cadenat (Bull. Inst. Francais Afrique Noire, vol. 21, ser. A., no. 1, 
pp. 361-385, figs. 1-26, 1959) described Antennarius (Fowlerichthys) 
senegalensis, A. (Triantennatus) delaisi, and A. (T.) occidentalis as 
new species from off the west coast of Africa. 

Golem Whitley 

Xenophrynichthys Schultz (synonym). 

The generic name Golem Whitley (type species, Antennarius 
cryptacanthus Weber), published May 8, 1957 (Proc. Roy. Zool. Soc. 
New South Wales, p. 70) has priority over Xenophrynichthys Schultz 
(Proc. U.S. Nat. Mus., vol. 107, no. 3383, p. 81, November 1, 1957), 
both with the same type species. 

Antennarius radiosus Ganuan 

Kanazawaichthys scutatus Schultz (synonym). 

In 1957 (op. cit., p. 63) I described Kanazawaichthys scutatus on 
prejuvenile specimens, characterized by having two pairs of enlarged 
bony plates on the head. Hubbs (Copeia, no. 4, pp. 282-285, 1958), 
with additional specimens and aided b}^ me in a restudy of still more 
specimens, concluded that K. scutatus is the prejuvenile of A. radiosus, 
an opinion with which I concur. 

Maul (Bocagiana, Mus. Mun. Funchal, no. 1, p. 15, 1959) records 
a large specimen of A. radiosus from Madeira, and Palmer (Ann. Mag. 
Nat. Hist., ser. 13, vol. 3, pp. 149-151, 1960) records a prejuvenile 
A. radiosus, 13 mm. in standard length, off the West Coast of Ireland, 
54°10' N., 12°10' W. ; thus, this western Atlantic species occasionally 
occurs in the eastern part of the Atlantic Ocean. 



178 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

Lophiocharon Whitley 

Plumantennatus Schultz (synonym). 

Lophiocharon caudimaculatus (Riippell) 

Antennarius asper Macleay (synonym). 

The main character I used for the separation of L. caudimaculatus 
(subgenus Lophiocharon) and L. asper (type of the subgenus Pluman- 
tennatus) was a supposed difference in the bait, a simple tentacle in 
the former and a plumelike bait in the latter. I now find the nature 
of the bait to be variable, from simple to plumelike, and, therefore, 
I synonymize Antennarius asper Macleay with L. caudimaculatus 
(Riippell) and the subgenus Plumantennatus Schultz (1957, p. 89) 
with Lophiocharon Wliitley. 

The usual color pattern of i. caudimaculatus is that figured by Schultz 
(1957, pis. 6, A, D, and 8, C) with or without an ocellate spot just 
behind the midbase of the soft dorsal fin; otherwise, the entire fish is 
brown-speclded or reticulated with brown. The chief distinguishing 
coloration, however, is in the caudal fin and consists of white or 
clear spots encircled with brown or blackish pigment. 

The following specimens were studied at the Department of Sys- 
tematic Biology, Stanford University (SU): SU 35781, Singapore 
Market, May 1937, A. W. Herre, two, 84 and 85 mm. standard length; 
30652, Singapore, A. W. Hcrre, two, 52 and 80 mm.; 20204 (holotype 
of A. lithinostomus Jordan and Richardson), Cuyo, Philippine Islands, 
R. C. McGregor, 84 mm.; 39498, Singapore, October 10, 1940, A. W. 
Herre, two, 45 and 83 mm.; 30651, Singapore, March 14, 1934, A. W. 
Herre, four, 39 to 81 mm.; 27872, Sandaken, British North Borneo, 
July 3, 1929, A. W. Herre, three, 65 to 87 mm.; 32730, Singapore 
Harbor, May 7, 1937, A. W. Herre, three, 53 to 68 mm. At the 
California Academy of Sciences (CAS) (without number or locality), 
five specimens measured 57 to 92 mm. 

Among the 26 specimens examined, I find that 6 — SU 30652 (2), 
32730 (3), and CNHM 47248 (1)— represent a black color phase 
described as follows: general background coloration black, tentacle 
barred, bait white; white saddle in front of first soft ray of dorsal fin 
reaching to level of eye; white blotches just behind corner of mouth, 
above base of pectoral fin, near tips of third dorsal spine, near tips 
of rays of caudal, pectoral, and posterior rays of both soft dorsal and 
anal fins; white bar between rear of bases of soft dorsal and anal fins, 
and another white bar across base of caudal fin raj^s; sometimes rear 
third of caudal fin white and posterior margins of pectoral, pelvic, 
anal, and dorsal fins white; a few circular white spots may, or may 
not, occur in black area on middle of caudal fin. 



NEW INDO-PACIFIC FROGFISHES SCHULTZ 179 

Antennarius phymatodes Bleeker 

Antennarius oligospilos Bleeker (synonym). 

In my review (1957), I recognized as distinct species A. phymatodes 
and A. oligospilos on the basis of characters supposedly shown in 
Blocker's illustrations. Recent correspondence with Dr. Boeseman, 
along with information published by Dr. de Beaufort (The Fishes of 
the Indo-Australian Archipelago, vol. 11, pp. 204-206, 1962), con- 
clusively show that A. phymatodes and A. oligospilos represent the 
same species. 

These conclusions are based on the very close similarity of the holo- 
types of the nominal species as follows: 

A. phymatodes Bleeker, RMNH 6285, holotype (illustrated in 
Bleeker's Atlas, plate 199, fig. 5, 1865), standard length 69 mm., 
total length 93 mm.; dorsal rays 111,12, last 2 branched; anal 7, 
all branched; pectorals 10-10, none branched; pelvics 1,5, last ray 
branched; caudal 9, all branched. 

A. oligospilos Bleeker, RMNH 6286, holotype, standard length 
63 mm., illustrated (slightly modified after the 68 mm. nontype 
smaller of two specimens in RMNH 25010) as plate 195, fig. 1, in 
Bleeker's Atlas; dorsal rays 111,12, last 2 rays branched; anal 7, all 
branched; pectoral 10-10, none branched; pelvics, 1,5, last ray 
branched; caudal 9, aU branched. 

Description of New Species 
Phrynelox lochites, new species 

Plate 1 

This new species of Phrynelox with short "bait" (about the same 
length as the second dorsal spine) from the Philippine Islands has as a 
cognate species its closest relative, P. scaber of the western tropical 
Atlantic Ocean. 

Holotype, SU 38194, Dapitan Bay, Mindanao Island, Philippines, 
August 1940, collector A. W. Herre, standard length 48.5 mm. Three 
paratypes, all with same locality data as holotype: SU [number un- 
available], two specimens, 33.5 and 40.0 mm.; and USNM 197325, 
39 mm. 

The following counts were made: dorsal soft rays 12, anal 7, 
branched caudal 4+5, in all specimens; pectoral rays 11-11 in holotype 
and in two paratypes, 10-10 in one paratype. 

Aleasurements made on the types are recorded in table 2. 

Bony part of first dorsal spine a little longer than the second dorsal 
spine, bearing at its tip the fleshy bait, consisting of two robust 
tentacles, each with numerous small papillae; skin just behind base of 
second dorsal spine naked ; third dorsal spine movable posteriorly, tip 



180 



PROCEEDINGS OF THE NATIONAL MUSEUM 



free, but bound down with skin anteriorly, and subequal in length to 
second spine ; soft dorsal rays all simple except last one or two, which 
are branched; only last pelvic ray divided; caudal rays branched; 
anal rays all divided; pectoral rays all simple; gill opening close to 
base of pectoral fin; caudal peduncle distinct but short, slightly deeper 
than long; skin thicldy covered with bifid to multifid prickles; scat- 
tered dermal cirri present. 

Color in alcohol: head, body, and fins with light tan background; 
everywhere, except on underside of head and belly, with black spots 
or elongate blotches, arranged somewhat in lines on sides of body; 
black spots on fins round, those on caudal fin forming 4 vertical rows. 

In the genus Phrynelox, this new species is a member of the sub- 
genus Phyrnelox, which is characterized by having the "bait" formed 
of bifid tentacles. In this subgenus the species fall into two groups: 
(1) those species with the bony part (illicium) of the first dorsal spine 
notably long, almost twice the length of the second dorsal spine, 
including P. striatus (Shaw) and P. melas (Bleeker) ; (2) those species 



Table 2. — Measurements made on three new species of frogfishes {recorded in 
thousandths of the standard length) 





Phrynelox lochites 


Antennarius 




indicus 


japonicus 


Characters 


Holotype 


Paratypes 


Holotype 


Holotype 


Standard length In mm. 
Greatest depth of body 


48.5 
495 


40.0 
625 


39.0 
590 


33.5 
532 


44.5 

548 


49 
530 


Length of bony part of 1st dorsal spine 
" " " " "2nd " 
" " ' 3rd " " 


165 
124 
124 


175 
150 
150 


162 
115 
103 


179 
164 
143 


135 
168 

207 


112 
143 

208 


Longest soft dorsal ray 
Longest (middle) caudal ray 
Head (snout to gill opening) 


206 
289 
474 


238 
337 

575 


205 
318 
640 


239 
338 

686 


265 
355 
524 


204 
335 
550 


Length of caudal peduncle (or distance be- 
tween vertical lines through caudal fin base 
and rear bases of anal-dorsal fins) 


103 


112 


103 


119 


115 


69 


Length of maxillaries 

Least depth of caudal peduncle 


217 
124 


250 
145 


205 
115 


209 
119 


236 
153 


245 
133 


Length of base of soft dorsal fln 
Eye diameter 
Interorbital space 


454 
41 
144 


550 
60 
132 


474 

72 

141 


538 
60 
134 


540 
56 
157 


517 

75 

106 



NEW INDO-PACIFIC FROGFISHES — SCHULTZ 181 

with the bony part of the bait from a little longer to about the same 
length as the second dorsal spine, including P. scaber (Cuvier) of the 
western Atlantic and the black color phase of this species, known 
formerly as "P. nuttingi Garman." 

All of the species of the first group that are known so far are from 
the western Pacific and Indian Oceans, whereas, in the second group, 
only P. lochites is found in the western Pacific, the remainder being 
found in the Atlantic. P. lochites differs from its closest relative in 
the Atlantic, P. scaber, in having the bait about as long as the eye 
diameter instead of its being two or more times longer. The only 
color difference noted is that on the caudal and anal fins, the spots 
being smaller and in more regular rows than those in P. scaber. 

The species is named lochites ("lochites," masculine, Greek, meaning 
"lying in wait for their prey") to refer to the habit of frogfishes. 

Antennarius japonicus, new species 

Plate 2 

Holotype, SU 26796, Sagami Bay, Japan, collector A. Owston, only 
known specimen, 49 mm. standard length. 

The following counts were made: dorsal soft rays 12, all simple or 
unbranched; anal 7, first 3 simple, last 4 branched; caudal 4 + 5, all 
branched; pectoral 10-10, all simple; pelvic rays 1,5, last branched. 

Bony part of first dorsal spine shorter than second dorsal spine, 
bearing a tuft of tentacles at the tip; skin just behind second dorsal 
spine naked; third dorsal spine movable and longer than second 
spine; gill opening close to base of pectoral fin; caudal peduncle 
distinct but short, deeper than long; skin thickly covered with bifid 
to multifid pricldes; scattered dermal cirri present. 

Color in alcohol: background coloration light gray with six round, 
brown-edged white areas; one on rear of head with two smaller round 
ones below, in front of pectoral base; another below space between 
third dorsal spine and origin of soft dorsal fin, and two on side of 
body; a large dark spot on body below bases of 8th to 10th rays of 
dorsal fin; color greatly faded but dorsal, anal, and caudal fins show 
traces of light brown spots; area above base of pectoral fin brownish, 
with light center. 

Remarks: This species traces through my key of the Antennariidae 
(1957, pp. 53-62) to section 35a, but it differs at this point by having 
the first dorsal spine shorter than the second dorsal spine. Also, this 
species differs from Antennarius pardalis (section 36a) by having 12 
dorsal rays, and from A. pardalis and A. bermudensis (section 36b) by 
having a distinctive color pattern of large round white spots. 

The new species differs from A. sarasa Tanaka (section 33b) in 
having all dorsal rays simple and 10 pectoral rays, whereas, in the 



182 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Japanese species, 8 of the dorsal rays are branched and there are 
13 pectoral rays. 

Another species recorded from Japan, A. nummifer (section 51b), 
is close to this new species, but it differs in having all of the anal rays 
divided and the last 2 or 3 dorsal rays divided at their tips. The 
number of fin rays in the two species agrees but the color pattern of 
A. nummifer does not show round white spots like those of 
A. japonicus. 

This species is named japonicus in reference to the country where 
the holotype was collected. 

Antennarius indicus, new species 

Plate 3 

Holotype, SU 40090, Vizagapatam, India, Dec. 25, 1940, collector 
A. W. Herre, only known specimen, 44.5 mm. standard length. 

The following counts were made: dorsal rays 12, last 2 branched; 
anal 7, all branched; caudal 4 + 5, all branched; pectoral 13-13, all 
simple; pelvic rays 1,5, last ray branched. 

Bony part of first dorsal spine shorter than second dorsal spine, 
bearing a tuft of tentacles at tip; skin just behind second dorsal spine 
naked; third dorsal spine movable and slightly longer than second 
spine; gill opening close to base of pectoral fin; caudal peduncle 
distinct, deeper than long; skin thickly covered with bifid to multifid 
prickles; scattered dermal cirri present. 

Color in alcohol: background coloration gray, with a dark spot 
in middle of side of body, belly plain light gray; anal and caudal 
fins with scattered dark spots; outer third of dorsal fin with 6 dark 
bars, and a larger dark spot between bases of 7th to 9th ray; dorsal 
surface of pectoral fin with dark spots except tips of rays white, under- 
side of pectoral uniformly light gray; body appears to have had other 
dark markings, now indistinct. 

Remarks: This species traces through my key of the Anten- 
nariidae (1957, pp. 53-62) to section 49b, but it differs from section 
50a (A. verrucosus) and 50b (A. altipinnis and A. pauciradiatus) 
by having 13 pectoral fin rays, instead of 9-11, a difference thought 
to be of significance in this family. Among the various species 
referable to the subgenus Antennarius, only two have as many 
pectoral rays as 12 or 13: A. sanguineus from the eastern Pacific 
and A. dromhus from Hawaii and Cocos Island. The caudal peduncle 
in both species is very short, its depth 2^ or 3 times its length, whereas, 
in A. indicus, the depth is 1% its length. The lower sides and belly 
of A. dromhus and A. sanguineus are dark-spotted, whereas A. indicus 
lacks spots ventrally. 

The new species is named indicus in reference to the country where 
the holotype was collected. 

U.S. GOVERNMENT PRINTING OFFlCEil964 



Proceedings of /^P^K 

the United States fp^^^ W 
National Museum ^^^^ 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 



Volume 116 19M Number 3501 



NEOTROPICAL MICROLEPIDOPTERA, V 

SYNOPSIS OF THE SPECIES OF THE GENUS PROEULIA 

FROM CENTRAL CHILE 

(LEPIDOPTERA : TORTRICIDAE)i 



By Nicholas S. Obraztsov 



The Neotropical species usually referred to the genera Eulior 
Hiibner, Tortrix Linne, and Cnephasia Curtis actually have little in 
common with these Holarctic genera. Some of these species have 
been relegated recently to the genera Argyrotaenia Stephens, Sub- 
argyrotaenia Obraztsov, Proeulia Clarke, Anopina Obraztsov, and 
others, but the generic position of many species still remains uncertain. 
Five new species from Central Chile described in this paper and four 
already known are assigned to the genus Proeulia. Additional infor- 
mation is included here concerning the previously described species. 

The genus Proeulia was established recently for two species from 
the Juan Fernandez Islands (Clarke, 1963). The characters of this 
genus, originally based on a study of its type, P. robinsoni (Aurivillius), 



I Prepared with the aid of a National Science Foundation Grant. Previous 
parts of this same series are: I and II, Clarke, 1962, Proc. U.S. Nat. Mus., vol. 
113, no. 3457, pp. 373-388; III, Clarke, 1964, ibid., vol. 115, no. 3480, pp. 61-84; 
IV, Duckworth, 1964, ibid., vol. 116, no. 3497, pp. 97-114. 

183 



731-763—64 



184 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

and P. griseiceps (Aurivillius), have been confirmed by the additional 
nine species from Central Chile examined by the present author. 
Only in the description of the wing venation are some modifications 
necessary. In the original diagnosis of Proeulia, the veins R and Mi of 
the hindwing were described as stalked, and they are similar in most of 
the species treated in the present paper. In the new species Proeulia 
triguetra, however, these veins are either variously long-stalked, or 
originating connate, or even slightly separate, although closely ap- 
proximated to each other in the basal portions. Also, veins M3 and 
Cui of the hindwing are either connate (as in P. robinsoni and P. 
griseiceps) or slightly separate at origin. In view of the observed 
individual variation of these characters in one and the same species, 
it seems to be expedient to extend the diagnosis of the genus to include 
all these modifications. 

The study of the nine additional species assigned to this genus has 
demonstrated some important specific modifications in the shapes of 
certain parts of the genitalia, as is seen from the descriptions and 
photographs in this paper. The most unusual is the process that 
projects from the ventral surface of the bursa copulatrix, exhibiting a 
greater reduction in Proeulia griseiceps than in P. robinsoni. In P. 
auraria (Clarke), P. chrysopteris (Butler), and P. apospasta, new 
species, this process is very short. It is quite possible that in some 
species not yet known this reduction may appear to be even more com- 
plete. In P. leonina (Butler) and P. aethalea, new species, the above 
process is especially well developed. The location of this process close 
to the inception of the ductus seminalis and its connection with the sur- 
rounding sclerotization supports the identification of this structure as 
a cestum, developed in the present case as an external protrusion. The 
area around the ostium oviductus generally is sclerotized somewhat in 
all species of Proeulia. The corresponding sclerite is reversely sub- 
cordate as a rule and probably represents a modification of the papillae 
genitales known in some other families of the Lepidoptera (Kusnezov, 
1916). This structure still is studied poorly in the Tortricidae and 
probably will be found in other genera of this family. 

It is a pleasure for the author to express his gratitude to Dr. J. F. 
Gates Clarke of the U.S. National Museum and Mr. J. D. Bradley of 
the British Museum (Natural History) for the materials placed at his 
disposal and to Dr. J. G. Rozen and Dr. F. H. Rindge of the American 
Museum of Natural History for providing the necessary working 
facilities. The work on this paper was done under the auspices of the 
National Science Foundation ; many of the specimens involved in the 
study were collected by Dr. Clarke during field investigations sup- 
ported by the National Science Foundation. 



SPECIES GENUS PROEULIA — OBRAZTSOV 185 

Key to the Species of Proeulia Jfrom Central Chile 

1. Forewing unicolorously ocherous with some few, little, scattered black 

dots 2 

Forewing not ocherous; if ocherous, more markings, expecially at dorsum, 
are present 3 

2. Hindwing pale ocherous, at base with a concolorous or slightly darker tuft of 

hairs; sacculus not protruding valva leonina (Butler) 

Hindwing shiny white, at base with a white tuft of hairs; sacculus protruding 
valva cneca, new species 

3. Forewing with two bands originating at costa and meeting each other at a 

point slightly remote from dorsum tenon tias (Meyrick) 

Forewing without such bands 4 

4. Dorsum of forewing with a large triangular spot connected with wing base by 

means of a band along dorsum; hindwing dark hazel grayish to dark lead 

gray triquetra, new species 

Dorsum without any defined spot; if such is present, not connected with 
wing base ; hindwing whitish or somewhat fuscous, never very dark . . 5 

5. Forewing dark fuscous, diffusely shaded, without defined markings. 

aethalea, new species 
Forewing of some other color; markings more or less distinct and consisting 
either of a dorsal spot or of some (at least incomplete) bands and/or reticula- 
tion, or all these markings present 6 

6. Forewing with a ferruginous blotch in basal portion; an oblique, white line 

closely basad from dorsal spot apospasta, new species 

Forewing without ferruginous blotch in basal portion and no white line basad 
from dorsal spot 7 

7. Sacculus protruding valva and ending with a free, sharp point. 

chrysopteris (Butler) 
Sacculus not protruding valva and not pointed at tip 8 

8. External margin of valva curved in lower portion and reaching end of sacculus 

vertically; shorter cornutus longer than one-half length of long cornuti. 

inconspicua, new species 

External margin of valva not perceptably changing its direction from apex to 

end of sacculus; shorter cornutus less than one-half length of long cornuti. 

auraria (Clarke) 

Proeulia leonina (Butler), new combination 

Plate 1 

Sciaphila leonina Butler, 1883, Trans. Ent. Soc. London, p. 69. 
Torlrix leonina, Meyrick, 1912, in Wagner, Lepidopterorum catalogue, pt. 10, 
p. 27; 1913, in Wytsman, Genera insectorum, fasc. 149, p. 29. 

Male genitalia: Uncus rather short; socii long, gradually and 
moderately dilated. Fultura superior narrow; fultura inferior high 
and rather narrow. Valva broad; sacculus moderate, pointed, not 
protruding valva. Aedeagus moderately thickened, with tip rounded; 
three long cornuti, one longer than remaining two. 

Female genitalia: Lamella antevaginalis large, deeply incised at 
middle, covering ostium bursae. Antrum large, subrect angular. 



186 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Corpus bursae elongate-ovate, with a large, lateral swelling bearing 
a well-developed cestum; cervix bursae broad and short. Sclerotiza- 
tion around ostium oviductus well developed. 

Tjrpes: Lectotype, male (genitalia on slide 8641), Valparaiso, 
Chile, November 1881 (T. Edmonds, 1882-107); lectoaUotype, female 
(genitalia on slide 8776), same data; British Museum (Natural 
History), 

Other specimens examined: One male and two females, Chile (T, 
Edmonds, Druce Collection, 1917-36); one female (genitalia on slide 
6711), Quillota, Valparaiso, Chile, 1886 (Paulson, 68384); British 
Museum (Natural History). 

Proeulia auraria (Clarke), new combination 

Plates 2, 3 
Eulia auraria Clarke, 1949, Acta Zool. Lilloana, vol. 7, p. 583, pi. 2, figs. 6-Cb 

The original description of this species was based on a series of six 
male specimens, all of them taken in Caj6n de Maipo, Santiago. 
Many additional specimens examined for the present paper originated 
from other localities, showing that the species is rather variable. 
The length of the fore wing varies from 7 to 12 mm. Most of the 
small specimens (the type series included) have the forewing markings 
reduced to an incomplete oblique band running from the middle of 
the costa toward the tornus and sending from slightly below its 
middle a short band directed toward the middle of the dorsum. These 
bands are generally underdeveloped, and only some of their elements 
are present in each specimen. In some specimens the dorsal portions 
of the above-mentioned bands form a more or less distinct dorsal 
triangle or a semicircular arch, some outlines of which are dilated and 
more intensively dark colored, forming blackish-brown spots. In addition, 
one more band may be present, running externally to the oblique 
costotornal band and parallel to it. Occasionally all or some of the 
bands are widely dilated, and/or some minute, brownish, or ferrugi- 
nous streaks at the costa and dorsum are present. The golden- 
ocherous ground color of the forewing is more or less mottled with 
brown or ferruginous, which causes development of a fine reticulation 
or separate spots or short lines. Rarely the two basal thirds of the 
forewing or even the entire forewing are overlaid with brown. Upper 
basal angle of the forewing commonly differs from the ground color 
of the wing and is brown, gray, or yellowish. The hindwing is white, 
ocherous white, or pale cinereous, in some specimens distinctly speckled 
with gray. 

Male genitalia: Uncus rather long, slightly bent; socii dilated 
externally. Fultura superior narrow; fultura inferior moderately 



SPECIES GENUS PROEULIA — OBRAZTSOV 187 

high. Valva elongate, gradually bent upward and slightly narrowed 
apicad; sacculus moderate, somewhat tapering apicad, not free 
pointed. Aedeagus rather stout; from two to four long cornuti, 
one short, slightly curved, less than half length of long cornuti. 

Female genitalia: Lamella antevaginalis band shaped, narrowed 
laterad; lamella postvaginahs semimembranous. Antrum not de- 
veloped. Corpus bursae somewhat ovate, scobinate; cestum shaped 
as a short, rotundate projection with a moderately sclerotized base; 
cervix bursae broad and short. Sclerotization around ostium ovi- 
ductus A-shaped. 

Types: Holotype, male (genitalia on slide), and three male para- 
types in USNM. One male paratype in the British Museum (Natural 
History). One male paratype (not seen by the present author) in the 
collection of F. Bourquin. All were collected in Caj6n de Maipo, 
Santiago, Chile, Jan. 12-20, 1948 (Don Tito Ramirez). 

Other specimens examined: One male, Chile Centro-Austral, Jan- 
uary-March 1898 (V. Izquerdo), in USNM. Five males (genitalia 
of two on slides 6694 and 6703), and two females (genitalia of one on 
slide 6710), Coquimbo, Chile, July 1-Oct. 19, 1883 (Walker, 3021- 
3024, 3000, 3205), and May 1884, bred from Aristolochia chilensis 
(Walker, 3234); two males (genitalia of one on slide 6760) and one 
female, Valparaiso, Chile, Mar. 22-Apr. 13, 1882, Sept. 30-Oct. 8, 
1883 (Walker, 2353, 2359, 3078); six males (genitaha of one on sHde 
6693), Quillota, Valparaiso, Chile, 1886 (Paulson, 68013-68018); 
the foregoing 15 specimens are deposited in the British Museum 
(Natural History). Five males (genitalia of one on slide, 21-Obr., 
1962), Guayacan, Santiago, Chile, 1100 m., Jan. 25-26, 1951, Oc- 
tober 1952 (L. E. Pena); two males (genitalia on slides, 1-Obr., 
1962, and 2-Obr., 1962), La Obra, Santiago, Chile, October 1952 
(L. E. Pena) ; one male (genitalia on slide, 22-Obr., 1962), El Principal, 
Chile,' November 1888 (V. Izquerdo); all eight specimens are in 
USNM. 

Remarks: The holotype and its genitalia were figured by Clarke 
(1949). The shape of the valvae became distorted slightly on the 
original photograph, and they look more narrow apicad than they 
really are. The holotype has three long cornuti, but this number 
varies in the species. In one of the males examined (slide, 1-Obr., 1962) 
there are only two long cornuti; another male (slide, 21-Obr., 1962) 
has four of them. The presence of a short, slightly bent comutus is 
apparently constant for the species. The variation of the number of 



1 Because the collector is deceased, there is probably no way of ascertaining 
exactly this location. Gazetteer (no. 6, Chile, Office of Geography, Department 
of the Interior, Washington, August 1955, p. 107) lists two names, one located at 
33.47 S. and 70.30 W., the other at 33.42 S. and 70.34 W. 



188 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

the cornuti and the forewing markings are independent from each 
other. 

Proeulia aethalea, new species 

Plate 4 

Antenna fuscous. Labial palpus pale brownish gray, speckled with 
dark fuscous. Head fuscous touched with gray. Thorax grayish 
brown. Abdomen brownish gray with a dark brown transverse line 
on caudal edge of eighth abdominal tergite. Legs ocherous. Fore- 
wing fuscous with indistinct, somewhat olive-brownish or gray- 
brownish markings apparently consisting of two diffuse, very broad 
transverse bands inclined basad; cilia dark brownish gray (as seen 
under magnification, with scales white checked). Underside of 
forewing grayish ocherous. Length of forewing 11 mm. Hindwing 
yellowish white, mottled with pale olive brown; an olive-brownish 
hair brush at base of veins A2 and A3; cilia whitish. 

Male genitalia : Uncus rather long ; socii dilated externally. Fultura 
superior dilated laterally; fultura inferior triangular, moderately high. 
Valva elongate with apex directed upward and external margin 
rather vertical, in lower portion; sacculus rather broad, moderately 
long, with tip slightly tapering and slightly protruding beyond lower 
margin of valva. Aedeagus thickened, with a long and broad lower 
distal portion and a sclerotized margin around orificium; cornuti 
arranged in two groups: upper of eleven long cornuti, and lower of 
three cornuti about half as long as upper ones. 

Female genitalia: Lamella antevaginalis shaped as a semicircular 
band dilated toward antrum; lamella postvaginalis weakly sclero- 
tized, forming dorsal surface of funnel of ostium bursae. Antrum 
large, membranous. Corpus bursae elongate, sclerotized, covered 
with stronger sclerotized, longitudinal lines and, in caudal portion, 
with fine scobination; cestum in form of a long, digitate projection 
arising from a large basal disc located cephalad from cervix bursae. 
Weak sclerotization around ostium oviductus. 

Types: Holotype, male (genitalia on slide, 17-Obr., 1962), La Obra, 
Santiago, Chile, October 1902 (L. E. Pena), USNM (type no. 66830). 
Allotype, female (genitalia on slide 6699), Valparaiso, Chile, Sept. 
30-Oct. 8, 1883 (Walker, 3080), British Museum (Natural History). 

Remarks: This species differs from the remaining known members 
of the genus in having very indistinct and poorly defined markings on 
the forewing. The shape of the aedeagus, the set of the cornuti, 
and the characters of the female genitalia are unique in the genus 
Proeulia. The name of this species is derived from the Greek 
ai9a\eos, meaning "smoky." 



SPECIES GENUS PROEULIA — OBRAZTSOV 189 

Proeulia triquetra, new species 

Plate 5 

Antenna more or less dark brown. Labial palpus ocherous, more 
or less strongly mixed with brown, dark brown, somewhat ferruginous 
brown, or gray brown, occasionally with slight grayish suffusion; on 
inside pale ocherous, at least at base. Head and thorax brownish 
ocherous, hazel brown, mahogany brown, or ferruginous brown, 
often somewhat grayish suffused; posterior scale tuft occasionally 
more intensively colored. Abdomen fuscous with occasional touch 
of pale gra3dsh or ocherous; ventral surface usually somewhat paler, 
occasionally grayish ocherous. Legs hazel grayish, on inside pale 
ocherous. Fore wing brownish ocherous, testaceous, bay, or of some 
approximate shade, with more or less distinct, brown reticulation; 
dorsum with a paler, whitish-ocherous, longitudinal streak slightly 
suffused with brownish ocherous, testaceous, or bay; it originates at 
wing base and is dilated into a large triangle before tornus; upper 
margin of this streak is outlined by a narrow, white line, occasionally 
missing in some places, but constant at inner side of dorsal triangle 
and acompanied there by a more or less distinct, brownish-ocherous 
to dark-gray line (or merely a shadow) from inside of triangle; 
occasionally dorsal streak and its triangle are strongly darkened, and 
indicated only by above-mentioned white Une along their upper 
margins; area, bordering dorsal streak and reaching discal cell, 
usually darker than adjacent ground; a brownish or blackish more 
or less distinct line originating at middle of costa, directed to, or to 
slightly below, apex of dorsal triangle, then turning toward wing 
apex; external portion of this line occasionally dilated, outlining 
poorly defined costal triangle before wing apex; some occasional, 
minute, blackish strigulae perpendicular to costa and/or dorsum; 
cilia concolorous with wing ground, occasionally dark grayish checked, 
or darker tornad, rarel}^ paler at tips; underside of forewing dark 
brownish gray, at margins brownish ocherous or testaceous ; occasion- 
ally entire central area somewhat lightened, grayish testaceous. 
Length of forewing 9-11 mm. Hindwing hazel grayish to lead gray; 
cilia concolorous or slightly paler, with a fine, more or less dark 
gray line. 

Male genitalia: Uncus rather long; socii strongly dilated and 
rounded. Fultura superior rather broad; fultura inferior moderately 
high. Valva elongate, directed obliquely upward; sacculus broad, 
short, not free pointed. Aedeagus thickened; three long cornuti, 
and a separate group of 11-12 very short ones. 

Female genitalia : Lamella antevaginalis bandlike, narrowed lateral- 
ly; lamella postvaginalis weakly sclerotized. Antrum wide and short. 



190 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Corpus bursae pyriform; scobination chiefly developed at its left side; 
cestum shaped as a rather narrow, digitate projection on a wealdy 
sclerotized base; cervix bursae broad, tubular. Sclerotization around 
ostium oviductus well developed. 

Types: Holotype, male (genitalia on slide, 7-Obr., 1962), and allo- 
type, female (genitalia on slide, 10-Obr., 1962), Chilian, Nuble, Chile, 
Nov. 10, 1961 (H. Lister). Paratypes: one female (genitalia on slide, 
9-Obr., 1962), San Ignatio, Nuble, Chile, February 1892 (V. Izquerdo); 
four males (genitalia of one on slide, 5-Obr., 1962), Concepcion, Chile, 
March 3, 16, and 30, 1961 (Trampas); one female, Araucania [Arauco], 
Chile, March 1, 1892 (V. Izquerdo); three females (genitalia of one 
on slide, 8-Obr., 1962), Chile (V. Izquerdo; Silva). All specimens are 
deposited in USNM (type no. 66831). 

Remarks: This is the only known Proeulia species with a broad 
dorsal streak on the forewing. In other species, the dorsal triangle 
is not connected by a streak with the wing base. The genitalia of 
both sexes of P. triquetra have many characters distinguishing it from 
other species of the genus. 

Proeulia inconspicua, new species 

Plate 6 (Figs, 4-6) 

Male: Antenna ocherous with brown annulation. Labial palpus 
ocherous with slight ferruginous tinge, on inside pale ocherous. Head 
pale ocherous. Thorax (badly damaged) ferruginous ocherous ; tip 
of tegula ocherous. Abdomen pale ocherous, slightly brownish 
dorsally. Legs ocherous. Forewing ocherous, irregularly, densely 
sprinkled with ferruginous; at middle of dorsum a short, broad, 
ferruginous brown streak inclined basad ; an inconspicuous, ferruginous 
fascia running from middle of costa to torn us; transverse rows of 
minute, ferruginous dots, in external wing portion ; some inconspicuous, 
minute, ferruginous costal streaks, darker of them located apically; a 
fine, ferruginous terminal line, gradually becoming obsolete tornad; 
cilia pale ferruginous ocherous, slightly darker at tornus ; underside of 
forewing ocherous, somewhat ferruginous costad, with transverse 
rows of inconspicuous, minute, brownish-ferruginous dots. Length 
of forewing 11 mm. Hindwing very pale fuscous white, almost white, 
sprinkled with gray; hair tuft on wing base pale gray; cilia concolorous 
with wing surface. 

Female: Unknown. 

Male genitalia: Uncus rather long; socii elongate, dilated at middle. 
Fultura superior moderately and equally broad; fultura inferior almost 
triangular, moderately high. Valva elongate, directed upward; its 
external margin vertical, in lower portion; sacculus broad, rounded 



PROC. U.S. NAT. MUS. VOL. 116 



OBRAZTSOV— PLATE 1 








i^'- 



\ 



5 



r 



'm 



1 



Plate 1. — Proeulia leornna (Butler). Lcctoljpc, male: 1, right wings (iinape reversed); 
2, \entral aspect of genitalia with aedeagiis removed; 3, lateral aspect of aedeagus. 
Lectoallotype, female: 4, ventral aspect of genitalia; 5, detail of bursa copulatrix. 
Female genitalia of other specimen (slide 6711): 6, \-cntral aspect of genitalia; 7, detail 
of bursa copulatrix. 



PROC. U.S. NAT. MUS. VOL. 116 



OBRAZTSOV — PLATE 2 






Plate 2 — Proeulia auraria (Clarke), males. Specimen from Guayacan, Jan. 25, 1951: 1, 
left wings. Genitalia (slide, 1-Obr., 1962): 2, ventral aspect with aedeagus removed; 3, 
lateral aspect of aedeagus; 4, same, great!}' enlarged. Aedeagus (slide, 21-Obr., 1962): 

5, lateral aspect greatly enlarged. Right wings of two specimens (images reversed): 

6, from El Principal, November 1888; 7, from Valparaiso, March 22-April 13, 1882. 



PROC. US NAT MUS. VOL. 116 



OBRAZTSOV— PLATE 3 






'J ■ ' 


\ 


V 


f 

4-^ 






k;. 


. n, 



Plate 3. — Proeulia auraria (Clarke). Left wings of two spccinuiiN: 1, male from Quillota, 
1886; 2, female from Coquimbo, July 1-Oct. 19, 1883. Female genitalia (slide 6710): 
3, ventral aspect; 4, detail of bursa copulatrix. 



PROC. U.S. NAT MUS. VOL 116 



OBRAZTSOV— PLATE 4 




Plate 4. — Proeulia aelhalea, new species. liolotype, male: 1, left wings; 2, ventral aspect 
of geniialia with aedcagus reni()\ed; .\ lateral aspect of aedeagiis. Allotype, female, 
genitalia: 4, ventral aspect; 5, detail of bursa copulalrix. 



PROC. US NAT. MUS. VOL. 116 



OBRAZTSOV - PLATE 5 




Plate 5. — Proeulia Iriquelra, new species. Holotype, male: 1, left wings; 2, ventral aspect 
of genitalia with aedeagus removed; 3, lateral aspect of aedeagus; 4, detail of aedeagus. 
Allotype, female: 5, left wings; 6, ventral aspect of genitalia; 7, detail of bursa copulatrix. 



PROC. U.S. NAT. MUS VOL, 116 



OBRAZTSOV — PLATE 6 




Plate 6. — Proeulia apospasta, new species, holotype, female: 1, right wings (image reversed); 
2, ventral aspect of genitalia; 3, detail of bursa copulatrix. P. inconspicua, new species, 
holotype, male: 4, left wings; 5, ventral aspect of genitalia with aedeagus removed; 6, 
lateral aspect of aedeagus. 



PROC. US. NAT. MUS VOL. 116 



OBRAZTSOV — PLATE 7 




Pl.\te l.— Proeulta tenontias (Meyrick), female (genitalia on slide, 19-Obr., 1962): 1, left 
wings; 2, ventral aspect of genitalia; 3, detail of ostium bursae; 4, cestum. P. cneca, 
new species, holotype, male: 5, left wings; 6, ventral aspect of genitalia with aedeagus 
removed; 7, lateral aspect of aedeagus. 



PROC. U.S. NAT. MUS. VOL. 116 



OBRAZTSOV — PLATE 




Platb; 8.— Prot'itliii chryjopte'ris (Butler), males Ilolot}"pe: 1, left wings; 2, ventral asp- L 
of genitalia with aedeagus removed; 3, lateral aspect of aedeagus; 4, detail of aedeagus. 
Specimen from Chile Centro-Austral (genitalia on slide, 11-Obr., 1962): S, left wings; 6, 
ventral aspect of genitalia with aedeagus removed; 7, lateral aspect of aedeagus; 8, detail 
of aedeagus. Specimen from Chile (genitalia on slide, 16-Obr., 1962): 9, right wings 
(image reversed). 



PROC U.S. NAT. MUS. VOu. 116 



OBRAZTSOV — PLATE 9 




Plate S.—Proeulia chrysopteris (Butler), females. Specimen from Chile Centro-Austral 
(genitalia on slide, 13-Obr., 1962): 1, left wings; 2, ventral aspect of genitalia; 3, detail of 
bursa copulatrix. Specimen of Guayacan: 4, left wings. Specimen from Santiago: 5, 
right wings (image reversed). 



SPECIES GENUS PROEULIA — OBRAZTSOV 191 

externally. Aedeagus rather thick, with lower tip acute and upper 
with a slightly elevated carina; two long cornuti, one short, slightly 
longer than one-half length of long ones. 

Type: Holotype, male (genitalia on slide, 3-Obr., 1962), La Obra, 
Santiago, Chile, October 1952 (L. E. Peiia), USNM (type no. 
66832). 

Remarks: Somewhat similar to Proeulia apospasta, new species, 
but the markings of the forewing are more indistinct; no distinct 
triangular spot on the dorsum of forewing. Very typical of the 
species is a carina in the upper distal portion of the aedeagus. 

Proeulia apospasta, new species 

Plate 6 (Figs. 1-3) 

Female: Antenna ocherous, sUghtly brownish annulated. Labial 
palpus ocherous, mixed with ferruginous, paler from inside. Head, 
thorax, and abdomen concolorous with labial palpus. Legs pale 
ferruginous ocherous. Forewing ocherous, slightly ferruginous 
ocherous in external portion, and strongly so colored at wing base; 
a broad, ferruginous, longitudinal blotch, originating at wing base, 
tapering basad and externad, and not reaching end of discal cell; 
a narrow, oblique, brownish-ferruginous streak from lower angle of 
discal cell to middle of dorsum; a narrow, whitish line separating 
this streak from external, oblique edge of above-mentioned ferruginous 
blotch, and connected to pale ocherous, dorsal area below this blotch; 
a narrow, oblique, slightly undulate, brownish-ferruginous band 
shaded externally by ferruginous, running costotornad from about 
middle of costa but not reaching tornus, and with its middle touching 
upper end of above-mentioned brownish-ferruginous oblique streak 
almost under right angle; a triangular, pale ocherous dorsal spot 
separated by that streak and lower portion of mentioned costotornal 
band; some indistinct, pale ferruginous, minute streaks in costal 
area, and fine, concolorous dots and lines in area externad from 
costotornal band; a minute, brownish-ferruginous dot at dorsum 
before tornus; cilia pale ferruginous, becoming pale ocherous tornad; 
underside of forewing pale ferruginous, abruptly becoming darker 
before a pale ocherous, subterminal line. Length of forewing 10 mm. 
Hindwing light fuscous with slightly brassy hue; a concolorous tuft 
of hairs at base of veins A2 and A3; cilia whitish with a fine, greyish 
basal line. 

Male: Unknown. 

Female genitalia: Lamella antevaginalis band-shaped, narrowed 
laterally; lamella postvaguialis weakly sclerotized. No separate 
antrum. Corpus bursae rotund ate-pyriform, in most of caudal por- 



192 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

tion scobinate; cestum located in cervix portion and consisting of a 
very short, rotundate projection on a bilobate, sclerotized base; cervix 
bursae rather broad. Sclerotization around otsiiim oviductus well 
developed. 

Type: Holotype, female (genitalia on slide, 18-Obr,, 1962), Concep- 
ci6n, Chile, October 1902 (E. C. Reed), USNM (type no. 66833). 

Remarks: Somewhat similar to Proeulia chrysopteris (Butler), but 
differing from it in some details of the markings of the forewing, 
especially in the presence of a ferruginous blotch in the basal portion 
of the wing, a whitish line basad from the brownish outline of the dor- 
sal triangular spot, and in having the hindwing unicolorous. The 
genitalic differences of P. apospasta consist of a flat sinus vaginalis, 
absence of the antrum, and especially of a bilobate base of the cestum, 
known only in this Proeulia species. The name of the species is 
derived from the Greek aTrosTrasroj, meaning "separated." 

Proeulia tenontias (Meyrick), new combination 

Plate 7 (Figs. 1-4) 

Cnephasia tenontias Meyrick, 1912, Trans. Ent. Soc. London, 1911, p. 685; 1912, 
in Wagner, Lepidopterorum catalogus, pt. 10, p. 49; 1913, in Wytsman, 
Genera insectorum, fasc. 149, p. 47.— Clarke, 1955, Catalogue of the type 
specimens of Microlepidoptera in the British Museum described by Edward 
Meyrick, vol. 1, p. 304, 

Eulia tenontias, Clarke, 1958, Catalogue of the type specimens of Microlepidoptera 
in the British Museum described by Edward Meyrick, vol. 3, p. 139, pi. 69, 
figs. 4-4b. 

Male genitalia: Uncus moderately long; socii short. Fultura 
superior moderately broad; fultura inferior rather low. Valva with 
apex directed straight upward; its external margin vertical, in lower 
portion slightly incurved ; sacculus narrow, long, free pointed. Aedea- 
gus moderately thickened, slightly bent; four inequally sized cornuti. 

Female genitalia: Lamella antevaginalis band shaped with lateral 
portions directed caudad and outlining ostium bursae like a bracket; 
lamella postvaginalis membranous, not defined. Antrum large, sub- 
rectangular. Corpus bursae irregularly subovate ; scobination occupy- 
ing most of surface; cestum shaped as a digitate, slightly curved 
projection on a broad, swollen base; cervLx bursae tubular, distinctly 
narrower than corpus bursae. 

Type: Holotype, male (genitalia on slide, 6344, J.F.G.C.), Chile 
("R. .05"); deposited in the British Museum (Natural History). 

Other specimens examined : Two females, Chile (V. Izquerdo) ; one 
female (genitalia on slide, 19-Obr., 1962), Chile Centro-Austral, 
January-March 1898 (V. Izquerdo); one female (genitalia on slide, 



SPECIES GENTJS PROEULIA — OBRAZTSOV 193 

20-Obr., 1962), Quilpue, Valparaiso, Chile, November 1897 (V. 
Izquerdo) ; all specimens are deposited in USNM. 

Remarks: This species has hitherto been known as a unique male 
specimen. The holotype of P. tenontias and its genitalia were figured 
by Clarke (1958). The female specimens examined by the present 
author are very similar to the holotype, and there is no doubt that 
they are conspecific with it. 

Proeulia cneca, new species 

Plate 7 (Figs. 5-7) 

Male: Antenna grayish ocherous, in basal portion with a slight 
ferruginous tinge. Labial palpus brownish gray, on outer surface; 
separate scales dark gray basally, and whitish ocherous at tips; 
inner surface white ocherous. Head concolorous with outer surface 
of labial palpus, differing in color of separate scales, brownish gray 
at their middles, and whitish ocherous basally and at tips. Thorax 
brown with some scales long, orange ocherous; tegula orange ocherous. 
Abdomen shghtly paler than thorax. Legs ocherous. Forewing 
ocherous with slight silvery shine and some areas covered with yellow 
scales especially dense in basal third of wing, along discal cell, costal 
and dorsal portions, and on external veins; minute, blackish dots on 
disc and in external wing portion; ciUa pale ocherous, here and there 
shghtly ferruginous, especially at tornus; underside of forewing 
ferruginous ocherous, whitish ocherous externally and dorsally. 
Length of forewing 11 mm. Hind wing white; basal tuft of hairs 
white. 

Female: Unknown. 

Male genitalia: Uncus rather long; socii long, almost equally 
broad. Fultura superior very narrow at middle, dilated and strongly 
sclerotized laterally; fultura inferior high. Valva elongate, not turned 
upward ; its external margin rather short ; sacculus long with a pointed 
tip protruding far beyond valva. Aedeagus moderately tliickened, 
with tip rotundate; one rather thick and long cornutus with infundibu- 
liform envelope at base. 

Type: Holotype, male (genitalia on shde, 23-Obr., 1962), Guayacdn, 
Santiago, Chile, 1100 m., October 1952 (L. E. Pena), USNM (type no. 
66834). 

Remarks: This species is very similar to Proeulia leonina (Butler) 
and differs from it in having no black dots on the discocellulars of the 
forewing and in having the hind^^dng shiny white. The genitalia are 
distinct in the two species. The name of this species is derived from 
the Greek kvtikos, meaning "pale yellow." 



194 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Proeulia chrysopteris (Butler), new combination 

Plates 8, 9 

Tortrix chrysopteris Butler, 1883, Trans. Ent. Soc. London, p. 69. 
Eulia chrysopteris, Meyrick, 1912, in Wagner, Lepidopterorum catalogus, pt. 10, 
p. 39; 1913, in Wytsman, Genera insectorum, fasc. 149, p. 39. 

Originally described from a single male specimen, this species has 
become known as being widel}^ distributed in Central ChUe. It is 
rather variable, having the forewings ocherous, golden ocherous, 
testaceous, or hessian brown with a more or less intensive, ferruginous- 
ocherous reticulation and/or incomplete, oblique rows of blackish 
or grayish dots in the apical wing portion and occasionally basad from 
it. The upper basal angle of the forewing generally stands out as a 
triangle of a different color from the remaining wing surface and is 
gray or whitish, concolorous with the thorax. On the dorsum, slightly 
before tornus, there is a large, more or less distinct, white to grayish- 
yellow or brownish-yellow triangular spot rather broadly outlined by 
brown and checked by short, gray, or brownish streaks along the 
dorsum. A slight, ferruginous-ocherous line sometimes connects the 
apex of this triangle with the middle of costa. The length of the fore- 
wing is from 10 to 13 mm. Hindwing whitish yellow to ocherous, 
becoming gray basad in most specimens. The head is generally 
concolorous with the thorax. 

Male genitalia: Uncus moderately long; socii long, equally broad. 
Fultura superior rather broad; fultura inferior high. Valva with apex 
directed upward; sacculus broad, sharp pointed at tip, protruding 
valva. Aedeagus moderately thickened; two or three long, variously 
thick cornuti; a minute, sclerotized plate in external portion of vesica. 

Female genitalia: Lamella antevaginahs bracket shaped, with lateral 
portions directed caudad and bearing angulate prominences toward 
ostium bursae. Antrum large, semirotundate. Corpus bursae pyri- 
form, densely scobinate; signum located close to its middle, rather 
small, situated on a swollen base; cervix bursae rather broad. 

Type: Holotype, male (genitalia on slide 8587), Chile ("82-107"); 
deposited in the British Museum (Natural History). 

Other specunens examined: One female, Chile (V. Izquerdo); one 
male and two females (genitalia on slides, 16-Obr., 13-Obr., and 14- 
Obr., 1962), Chile Centro-Austral, January-March 1898 (V. Izquerdo); 
m USNM. Two males (genitalia on slides 6695, 6698), Quillota, Val- 
paraiso, Chile, 1886 (Paulson, 68011, 68012), in the British Museum 
(Natural History). One female, Santiago, Chile, June 1955, reared 
from apricot fruit (G. Olalquiaga) ; one female (genitalia on slide, 4- 
Obr., 1962), Guayacan, Santiago, 1100 m., Jan. 25, 1951 (L. E. Pena); 



SPECIES GENUS PROEULIA — OBRAZTSOV 195 

one male (genitalia on slide, 11-Obr., 1962), Concepci6n, Chile, Octo- 
ber 1902 (E. C. Reed) ; in USNM. One male (genitalia on slide 6696), 
Talcahuano, Concepci6n, Chile, Feb. 20-March 5, 1884 (Walker, 
3197) ; in the British Musemn (Natural History). One male (genitalia 
on shde, 12-Obr., 1962), Araucania [Arauco], Chile, March 1, 1892 
(V. Izquerdo), USNM. One male (genitalia on shde 6697), Valdivia, 
Chile, 1901 (A, von Lossberg); in the British Museum (Natural His- 
tory). One female (genitalia on slide, 15-Obr., 1962), Petrohue, 
Llanquihue, Chile, March 11, 1959 (J. F. Gates Clarke), m USNM. 
Remarks: On the slide of the genitalia of the holotype in the British 
Museum (Natural History), the valvae are folded and the sacculi 
turned inward, crossing the inner surface of the valvae. The folding 
of the lower margin of the valva causes a slight change of shape in 
the latter, and the apex of the valva becomes less acute as demon- 
strated by other specimens placed in the same position as the holotype. 
The extreme shapes of the genitalia (as illustrated on plate 8, figs. 2 
and 6) do not represent structural differences but only indicate 
differences that result from the preparation of the genitalia. The 
number of the cornuti varies between two (in the holotype and on 
slide 6698) and three (on six additional slides). 



Literature Cited 

Clarke, J. F. Gates 

1949. Notes on South American "Tortricidae" (Lepidoptera) and descrip- 
tions of new species. Acta Zool. Lilloana, vol. 7, pp. 579-588, 
3 pis. 
1958. Catalogue of the type specimens of Microlepidoptera in the British 
Museum (Natural History) described by Edward Meyrick, vol. 
3, (2) +600 pp., 298 pis. 
1962. A new tortricid genus from South America. Proc. Biol. Soc. 
Washington, vol. 75, pp. 293-294. 
KUSNEZOV, N. J. 

1916. Contributions to the morphology of the genital apparatus in 
Lepidoptera: Some cases of gynandromorphism. Rev. Russe 
Ent., vol. 16, pp. 151-191. 



U.S. GOVERNMENT PRINTING OFFICE: 1964 



Proceedings of .^^^hi^^ 

the United States ((( 
National Museum ^^S^ 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1964 Number 3502 

NEOTROPICAL MICROLEPIDOPTERA, VI ^ 

GENERA ORSOTRICHA MEYRICK AND PALINORSA MEYRICK 
(GELECHIIDAE, OECOPHORIDAE) 



By J. F. Gates Clarke 



The moths of the genera Orsotricha and Palinorsa have been con- 
fused in collections for fifty years. Only the study of their types, 
plus additional material, permits clarification of their relationships 
at this time. 

Gelechiidae 

Orsotricha Meyrick 

Orsotricha Meyrick, 1914, Exotic Microlepidoptera, vol. 1, p. 269; 1922, in Wyts- 
man, Genera insectorum, fasc. 180, p. 45. — Fletcher, 1929, Mem. Dept. 
Agric. India, Ent. Ser., vol. 11, p. 156. — Gaede, in Bryk, 1938, Lepidop- 
terorum catalogus, part 88, p. 87. — Clarke, 1963, Catalogue of the type 
specimens of Microlepidoptera in the British Museum (Natural History) 
described by Edward Meyrick, vol. 4, p. 341, pi. 167, figs. 1-le. 

1 Prepared with the aid of a National Science Foundation Grant. Previous 
parts of this same series are: I and II, J. F. Gates Clarke, 1962, Froc. U.S. Nat. 
Mus., vol. 113, no. 3457, pp. 373-388; III, Clarke, 1964, ibid., vol. 115, no. 
3480, pp. 61-84; IV, Duckworth, 1964, ibid., vol. 116, no. 3497, pp. 97-114; 
V, Obraztsov, 1964, ibid., vol. 116, no. 3502, pp. 183-196. 

729 977— 64 1 197 



198 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Meyrick proposed Orsotricha for Topeutis venosa Butler, from Chile, 
and subsequently associated with it Pleurota literatella Busck and 
Orsotricha raptans Meyrick, and placed the genus in the family 
Oecophoridae. 

Butler's Topeutis venosa, the type of the genus Orsotricha, was 
described from a single female. Both the genitalia and the wing 
venation, particularly the crossvein between vein 8 and the cell of 
the hindwing, indicated at the time of the study of the Me3"rick types 
that the species belonged in the Gelechiidae. Recent study of a male, 
the second specimen known, confirms this relationship and I hereby 
transfer Orsotricha to the Gelechiidae. 





Figure 1. — Orsotricha venosa (Butler): a, lateral aspect of male genitalia with aedeagus 

removed; b, aedeagus. 



NEOTROPICAL MICROLEPIDOPTERA, VI — CLARKE 199 

Orsotricha venosa (Butler) 

Figure 1; Plate 1 (Fig. 1) 

Topeulis venosa Butler, 1883, Trans. Ent, Soc. London, 1883, p. 77. — Meyrick, 
1914, Exotic Microlepidoptera, vol. 1, p. 269; 1922, in Wytsman, Genera 
insectorum, fasc. 180, p. 45. — Fletcher, 1929, Mem. Dept. Agric. India, 
Ent. Ser., vol. 11, p. 156. — Gaede, in Bryk, 1938, Lepidopterorum cata- 
logus, part 88, p. 87. — Clarke, 1963, Catalogue of the type specimens of 
Microlepidoptera in the British Museum (Natural History) described by 
Edward Meyrick, vol. 4, p. 341, pi. 167, figs. 1-le. 

Type: British Museum (Natural History). 

Distribution: Chile. 

The genitalia of the male are here figured for the first time. 

The two species previously associated with 0. venosa belong to the 
family Oecophoridae and are treated below. Fortunately, a generic 
name is available for them. 

Oecophoridae 

PaUnorsa Meyrick 

Palinorsa Meyrick, 1924, Exotic Microlepidoptera, vol. 3, p. 99. — Fletcher, 1929, 
Mem. Dcpt. Agric. India, Ent. Ser., vol. 11, p. 1.59.— Gaede, in Bryk, 1938, 
Lepidopterorum catalogus, part 88, p. 87. 

Meyrick proposed Palinorsa for Pleurota literatclla Busck after he 
had assigned that species to Orsotricha. Despite the obvious close 
relationship between P. literatclla and 0. raptans, Meyrick left the 
latter species associated with 0. venosa (Butler). I now include four 
species in Palinorsa. 

Key to the Species of Palinorsa 

1. Head brown or brownish 2 

Head ocherous white zonaria Clarke, new species 

2. Metathorax clothed with red hairlike scales 3 

Metathorax without red hairlike scales raptans (Meyrick) 

3. Forewing with conspicuous dark longitudinal streak. . . literatclla (Busck) 
Forewing without conspicuous dark longitudinal streak. 

acritomorpha Clarke, new species 

Palinorsa literatella (Busck) 
Figure 2 

Pleurota literatella Busck, 1911, Proc. U.S. Nat. Mus., vol. 40, p. 205, pi. 8, fig. 12, 
Palinorsa literatella (Busck), Meyrick, 1924, Exotic Microlepidoptera, vol. 3, p. 99. 

Type: USNM. 

Distribution: French Guiana, Rio Maroni. 

Male genitalia: Harpe sub triangular; cucullus rounded with serrate 
ventral edge produced as a strong thornlike process; overlapping 



200 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

the ciicullar process a double-pointed, flattened, sclerotized plate, 
serrate on outer edge. Anellus composed of two irregularlj^ shaped, 
curved plates fused at base and each with a small laterodistal lobe. 
Aedeagus short, stout, terminating in a beaklike process. Transtilla 
membranous, indicated by a few minute granules. Vinculum a 
broad sclerotized band. Gnathos an oval spined knob. Socii mainly 
indicated by setae. Uncus moderately hood-shaped, deeply incised 
on posterior margin. 

Slide examined: cT, JFGC 10948. 

Female genitalia : Ostium irregular, broadest posteriorly, surround- 
ed by a raised spiculate lip. Ductus bursae very short; inception of 
ductus seminalis dorsal and opposite center of ostial opening. Bursa 
copulatrix oval, without signum. Ovipositor lobes leathery, com- 
pressed. Anterior apophyses absent, or at most indicated only by a 
lateral thickening of genital plate. 

Slide examined: 9, JFGC 11198. 

This is the first time that the genitalia of P. literatella have been 
figured, and the female is the first of any species of this genus that I 
have seen. 

Palinorsa raptans (Meyrick), new combination 

Orsotricha raptans Meyrick, 1920, Exotic Microlepidoptera, vol. 2, p. 366. — Clarke, 
1963, Catalogue of the type specimens of Microlepidoptera in the British 
Museum (Natural History) described by Edward Meyrick, vol. 4, p. 341, pi. 
167, figs. 2-2b. 

Type: British Museum (Natural History). 
Distribution: Peru, Rio Napo. 

Palinorsa zonaria, new species 

Figure 3 

Alar expanse 42 mm. 

Labial palpus white; second segment with narrow fuscous line dorso- 
basally; third segment with slight apical infuscation. Antenna sordid 
white. Head ocherous white with ring of reddish-brown scales 
posteriorly. Thorax pale orange yellow shaded with reddish brown 
anteriorly; metathorax clothed with long, coral -red hau'like scales. 
Forewing pale orange yellow with brown median longitudinal streak: 
costa very narrowly edged with white; a pale spot on each side of 
vein Ic opposite base of vein 2; cilia pale orange yellow except at apex, 
the latter same color as median streak. Hindwing semihyaline; costal 
third and cUia ocherous white; at base a few coral-red hairlike scales 
and cilia. Legs ocherous white. 

Male genitalia: Harpe broad; saccular and costal margins parallel, 
distally tapering to a blunt cucullus ; middle heavily sclerotized giving 
rise to a strong thornlike clasper directed toward sacculus. Anellus 



NEOTROPICAL MICROLEPIDOPTERA, VI — CLARKE 



201 




Figure 2.—Palinorsa literatella (Busck): a, ventral view of male genitalia with left harpe 
and aedeagus removed; h, aedeagus; c, ventral view of female genitalia. 



U-shaped with broad basal plate fused with base of harpe. Aedeagus 
short, stout, terminating in a broad hooked process articulating with 
anellus. Vinculum a narrow ring. Gnathos spined, U-shaped. 
Socii indicated by numerous setae. Uncus elongate, narrowly tri- 
angular, terminating in two points. 
Slide examined: JFGC 11110. 



202 



PROCEEDINGS OF THE NATIONAL MUSEUM 





Figure 3.- — Palinorsa zonaria Clarke, new species: a, ventral view of male genitalia with 
left harpe and aedeagus removed; b, aedeagus. 

Female genitalia: Unknown. 

Holotype: Bolivia(?), Monte Cristo (and an indecipherable word), 
Amazones. USNM 64993. 

Described from the holotype male. Although there is only a single 
male with indefinite locality, it is so distinct it is worth description. 
The locality label leaves much to be desired. It is impossible to 
ascertain exactly where "Monte Cristo" is located but the "Amazones" 
suggests that the locality is actually in Bolivia. If it were possible 
to read the indecipherable part of the label, a more definite attribution 
might be made. In addition to the "Monte Cristo" label there is 
another in Busck's handwriting: "Orsotricha raptans Meyr.?" 

All of the species of this genus are closely related but zonaria 
appears to be nearest to literatella. It is distinguished easily from 
the latter by its ocherous-white head. 



NEOTROPICAL MICROLEPIDOPTERA, VI — CLARKE 



203 




Figure 4. — Palinorsa acritomorpha Clarke, new species: a, ventral view of male genitalia 
with left harpe and aedeagus removed; h, aedeagus. 

Palinorsa acritomorpha, new species 

FiGUHE 4; Plate 1 (Fig. 2) 

Alar expanse 33-36 mm. 

Labial palpus with second segment white ventrobasally, brown on 
outer side; on second segment a coral red median line on distal half 
on inner surface; third segment white. Antenna light broNXTi. Head 
cinnamon; face white. Thorax cinnamon brown with a broad ocher- 
ous-buff median longitudinal band; metathorax clothed with long coral- 
red hairlike scales. Forewing cinnamon; extreme costa, before apex, 
whitish; in cell at one-third, a small fuscous spot; at end of cell an 
ill-defmed transverse fuscous bar; on each side of vein Ic, opposite 
base of vein 2, a buff spot narrowly edged with fuscous; dorsum and 
termen narrowly fuscous; terminal cilia fuscous. Hindwing semi- 
hyaline whitish ; cilia sordid white, except around base, coral red. Legs 
white shaded with cinnamon and fuscous. Abdomen roseate dorsally 
with median longitudinal row of confluent cinnamon spots. 

Male genitalia: Harpe elongate ovate; cucuUar edge serrate termi- 
nating in a long spinous process; from outer third a large spine 



204 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

directed toward sacculus. Anellus V-shaped with prominent lateral 
lobes. Aedeagus short, stout, terminating in a beaklike process. 
Vinculmn a broad sclerotized band. Transtilla indicated by sparsely 
setaceous membrane. Gnathos a finely spined oval knob. Socii 
indicated by a few setae. Uncus elongate, incised posteriorly. 

Slides examined: 2crc?, JFGC 10946, 11109. 

Female genitalia: Unknown. 

Holotype: Peru, Tingo Maria (Nov. 24, 1949, H. A. Allard). USNM 
64992. 

Described from the type male and one male paratype with iden- 
tical data. 

The two species literatella and acritomorpho are similar but 
acriiomorpha lacks the dark longitudinal streak of the forewing 
found in literatella. 

All of the species of Palinorsa are related closely and one wonders 
if one is dealing with a "cline" or incipient species. All males possess 
a short, stout aedeagus terminating in a beaklike process, and all 
specimens have a pan' of small pale spots opposite the base of vein 
2 of forewing. The genitalia appear to present substantial char- 
acters for the separation of the species, but the paucity of material is 
a serious handicap to proper interpretation. 

Acknowledgments 

The photographs for the paper were made by Mr. Jack Scott, staff 
photographer. The drawings were made by Mr. Andre Pizzini. 



us. GOVERNMENT PRINTING OFFICE:1964 



PROC. U.S NAT. MUS. VOL. 116 



CLARKE— PLATE 1 




Figure 1. — Orsolricha vrnosa (Butler): lefl wings. 




Figure 2. — Palinorsa acriiomorpha Clarke, iu_\v sj'ccies: left wings. 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1965 Number 3503 

CONTRIBUTIONS TO THE KNOWLEDGE OF 

THE HEMEROBIIDAE OF WESTERN NORTH AMERICA 

(NEUROPTERA) 



By Waro Nakahara 



Introduction 



Dr. Charles P. Alexander [A] amassed a fine collection of the 
Hemerobiidae during his sixteen entomological expeditions to various 
parts of western North America, and this, together with the addi- 
tional material of nearly similar magnitude belonging to the United 
States National Museum [USNM], forms the basis of the present 
paper. Forty species recognized in the entire material are docu- 
mented, including five species that are described as new and another 
species that also may be new but is left unnamed for the present. 

The geographical area covered is west of the 100th meridian in the 
United States, British Columbia, Alberta, and Yukon in Canada, and 
Alaska. Localities by states and provinces not previously recorded 
are marked with asterisks. The reference to the original description 
is given for each species and genus; the complete synonymy is to be 
found in Carpenter (1940). 

Carpenter recognized 50 species of the Hemerobiidae as occurring 
in the entire Nearctic region. Later, Gurney (1948) and Parfin (1956) 
each added two species, and Nakahara (1960), one more. The addi- 

205 



206 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

tions made by the present study bring the total of the named species 
to 60. It is notable that as many as 57 of tliis number are known to 
occur in the area defined above. Western North America is thus a 
veritable treasure house of the Hemerobiidae, and it seems likely 
that further exploration may bring to light more new species, especially 
of Sympherobius, which obviously is collected inadequately, and of 
Kimminsia, which seems exceptionally rich in the area. 

My cordial thanks are due to Dr. Charles P. Alexander and Dr. 
Marion E. Smith for their kindness in submitting the Alexander 
collection to me for study. The major part of this collection has been 
returned to the Entomological Department, University of Massa- 
chusetts, Amherst, Mass., except the type specimens, which have 
been sent to the United States National Museum at the suggestion of 
Dr. Alexander. The examination of the United States National 
Museum material was made possible tlirough the kindness of Dr. 
Oliver S. Flint, Jr., and Dr. Ashley B. Gurney, to whom I wish to 
express my sincere appreciation. The whole of this material has been 
returned to the Museum. 

Family Hemerobiidae 

Subfamily Hemerobiinae ^ 

Genus Sympherobius Banks 

Sympherobius Banks, Proc. Ent. Soc. Washington, vol. 6, p. 209, 1904. 

The material before me contains ten species of this genus, including 
two that are new, while the following six species previously recorded 
are missing: S. umbratus Banks, S. arizonicus Banks, S. pictus 
(Banks), S. limbus Carpenter, ^S. similis Carpenter, and S. distinctus 
Carpenter, Most of these six species are known only from a unique 
type or at most from a few specimens. 

Sympherobius californicus Banks 

Sympherobius californicus Banks, Trans. Amer. Ent. Soc, vol. 37, p. 346, 1911. 

California: Alameda Co., Marin Co., San Jose, Benicia [USNM]. 
*Oregon: in Blue Mts. [USNM]. 

Sympherobius bifasciatus Banks 

Sympherobius bifasciatus Banks, Trans. Araer. Ent. Soc, vol. 37, p. 347, 1911. 

*Utah: Logan [USNM]. 



1 According to my studies on genitalia characters (1960), the family Hemero- 
biidae is divisible into two subfamilies, Hemerobiinae and Notiobiellinae, with all 
the Nearctic genera belonging to the former subfamily. 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 207 

Sympherobius angustus (Banks) 

Hemerohius angustus Banks, Trans. Anier. Ent. Soc, vol. 30, p. 102, 1904. 

♦Nevada: Mt. Charleston near Las Vegas [A]. *California: 
Yosemite Park [A]. Washington: Mt. Rainier [A]. Utah: Logan 
[USNM]. Colorado: Cheyenne Canyon [USNM]; Bierstadt Lake in 
Rocky Mountain National Park [A]. New Mexico: Las Vegas [USNM]. 

The head of this species was described by Carpenter as "dark 
brown, almost black." In many specimens, especially of small size, 
it is much lighter, often yellowish, with a large dark patch on frons 
below the antennae. I have dissected the male genitalia of a few 
specimens with the yellowish head and satisfied myself that they are 
not separable from S. angustus. 

Sympherobius stangei Nakahara 

Sympherobius stangei Nakahara, Mushi, vol. 34, p. 16, 1960. 

♦Colorado: Nymph Lake, Rocky Mountain National Park, 1 
female [A]. 

The specimen completely agrees with the unique type of S. stangei, 
except that it is slightly smaller (length of forewing 6 mm. against 
7 mm, in the latter). The original description is quoted here for the 
benefit of those to whom it may not be easily accessible: 

Head yellowish bro-mi, clypeus darker, palpi fuscous black, antennae fuscous 
black with paler basal joint. Forewing rather narrow, but much less so than 
in angustus, and fully rounded apically. Membrane colorless, broadly fuscous 
black along apical to outer margin; all veins behind Ri, except basal part of Cu2, 
distal part of first anal and basal half of second anal, broadly marked with fuscous 
black; cells thus strongly marked out are clear-colorless, not containing any spot. 
Hindwing less strongly margined with fuscous, veins dark but unmarked. 

Two branches to Rs in forewing, with radial crossvein between R4+5 and Ri be- 
fore the origin of R2+3; first fork of Cui distal to crossvein m-cu. 

Length of forewing, 7 mm; width 2.5 mm. 

Holotype: specimen lacking abdomen. Barton Flats, San Bernardino Co., 
California, July 22, 1953 (Lionel A. Stange). 

This is a large and beautifully marked species, perhaps related to occidenialis 
(Fitch). The striking markings and venational characters of forewing alone 
may be sufficient for the recognition of this new species. 

Sympherobius brunneus, new species 

Figure 1; Plate 1 (Figure 1) 

Holotype cf , Miami Ranger Station (elevation 5000 ft.), Mariposa 
Co., Cahf., July 5, 1945. Paratopotype cf, July 1, 1954. Both 
collected by H. P. Chandler. Right forewing (dry) and dissected 
parts of genitalia (in balsam) of holotype mounted on two slides. 
Paratopotype is without left forewing; terminal abdominal segments 
in glycerol in small vial on the same pin. USNM type 66174. 



208 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Face yellow, vertex slightly brownish; palpi dark brown; antennae 
dark brown with yellow fii'st joint. Pronotum yellow medially, 
narrowly brown on sides; meso- and matanotum yellow with light- 
brown scuta. Legs pale yellow. Abdomen light brown. 

Fore wing elongate oval with rounded apex, 5 mm. in length and 
2 mm. in width; costal area narrow; membrane nearly uniformly 
tinted with brown, without maculation; venation dark brown, longi- 
tudinal veins sparsely streaked with hyahne. A short radial cross- 
vein before origin of R2+3; inner gradates disjointed in middle, the 






c "* e 

Figure 1. — Sympherobius brunneus, new species: a, terminal abdominal segments, 
lateral view; b, the same, dorsal views; c, anal plate, internal lateral views; d, tenth 
sternite, lateral and slightly dorsal view; e, parameres, dorsal (above) and lateral view 
(below). 

upper two and lower two each on a straight line; Cui forked beyond 
crossvein m-cu. Hindwing membrane hyaline, tinted with brownish 
only in costal area, especially in pterostigmatic region. 

Male genitalia: Anal plate with four distal processes, resembling 
those of S. angustus, the first process from top (in lateral view) fairly 
long, the second the longest and most stout, the third the shortest, 
and the fom-th the second longest. Tenth sternite with very broad 
lateral "wing," which in lateral view appears roughly triangular with 
pointed distal apex. Parameres with long fused part, distal lobes 
parted at apex, forming an acutely narrow median incision; lateral 
flaps broader anteriorly than posteriorly. 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 209 

The brown-tinted membrane of forewing without maculation renders 
this species unique among the Nearctic Sympherobius. In order to 
include this species, Carpenter's key may be modified by introducing 
a new couplet 1' after couplet 1 (forewing with radial cross vein): 

r. Forewing membrane practically uniformly tinted with brown, without 

markings S. brunneus 

Forewing membrane with brown or grey markings . Carpenter's couplet 2 

Sympherobius barberi (Banks) 

Hemerobius barberi Banks, Proc. Ent. Soc. Washington, vol. 5, p. 241, 1903. 

Arizona: Williams, paratype no. 6798 [USNM]. Utah: Provo 
[USNM]. Texas: Kerrville and San Antonio [USNM]. 

Sympherobius perparvus (McLachlan) 

Hemerobius perparvus McLachlan, Ent. Monthly Mag., vol. 6, p. 22, 1869. 

*Utah: Tooele Co. [USNM]. California: Redding and Red Bluff 
[USNM]. Texas: Kerrville and Rankin [USNM]. 

Sym,pherobius beameri Gurney 

Sympherobius beameri Gurney, Ann. Ent. Soc. Amer., vol. 41, p. 220, 1948. 
California: Rosamond, paratype no. 58600 [USNM]. 

Sympherobius killingtoni Carpenter 

Sympherobius killingtoni Carpenter, Proc. Amer. Acad. Arts Sci., vol. 74, p. 238, 

1940. 

Arizona: Williams [USNM]. New Mexico: Las Vegas, paratype 
no. 55224 [USNM]. 

Sympherobius texanus^ new species 

Figure 2; Plate 1 (Figure 2) 

Holotype d": Kerrville, Texas, May 1954. Allotopotype 9, and 1 
paratopotype (without abdomen): June 1954. All collected by L. J. 
Bottimer. Right forewing (dry) and dissected parts of genitalia (in 
balsam) of holotype mounted on two slides. USNM type 66175. 

Face yellow, vertex somewhat more brownish. Antennae brown, 
darker toward apex. Pronotum pale brownish, with faint indication 
of yellowish median streak; meso- and metanotum yellow, with a 
large brownish patch on each side of metanotum. Abdomen yellow- 
ish, darker toward apex. 

Forewdng slender, 3 mm. m length, 1.25 mm. m width, but broadly 
rounded beyond middle, and hindmargin nearly straight, not curved out 
at the region of cubital forks; costal area very narrow. Membrane 
predominantly pale brown because of the more or less diffuse macula- 



210 



PROCEEDINGS OF THE NATIONAL MUSEUM 



tions, without any distinct blotch; venation pale brown, only outer 
gradates slightly and inner gradates strongly marked with bro\vn, the 
latter forming the only conspicuous marking; apical and outer 
margins very narrowly dark browTi w4th many pale interruptions. 
Radial crossvein absent. 




Figure 2. — Sympherobius texanus, new species: a, terminal abdominal segments, lateral 
viewj b, the same, dorsal view; c, anal plate, internal-lateral view; d, parameres, 
lateral view; e, tenth sternite, dorsal view, with aedeagus bent posteriorly. 

Male genitalia: Anal plate wdth ventroposterior angle slightly 
produced in lateral view; process single, fairly long, and straight. 
Tenth sternite with ventroproximal part of lateral "wing" expanded 
into a very large roundish flap. Parameres with the expanded distal 
part rounded on margin, without projecting middle arm. 

This species will fit into a modification of couplet 9 (species without 
radial crossvein) in Carpenter's key: 

9. Forewing membrane predominantly brown 9' 

Fore wing membrane predominantly hj^aline . to 10 of Carpenter's key 
9'. Forewing narrowed toward apex and hindmargin rounded out; larger 

species: forewing 4.5 mm. long S. arizoniciis Banks 

Forewing broadly rounded toward apex and hindmargin straight; 

smaller species: forewing 3 mm. long ... S. texanus, new species 

Texanus belongs to Gurney's "perparms group," which is charac- 
terized by the anal plate of the male bearing a single process and which 
includes S.perparvus, S. beameri, and S. killingtoni. Of these the latter 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 211 

may be the closest, but S. texanus can be distinguished readily by the 
very much smaller size and the brownish forewdng without blotchy 
markings. 

Genus Pseudomicromus Kriiger 

Pseudomicromus Kriiger, Stettin Ent. Zeit., vol. 83, p. 172, 1922. 

This genus, originally raised on the basis of ambiguous venational 
characters, has been reinstated by genitalic studies (Nakahara, 1960, 
p. 30). It is separated from other genera of the Micromus group by 
the paired aedeagus and by the absence of supraedeagal plate in the 
tenth sternite in the male. The material examined contained all the 
species that hitherto have been recorded. 

Pseudomicromus angulatus (Stephens) 

Hemerobius angulatus Stephens, Illustr. British Ins., vol. 6, p. 106, 1836. 

South Dakota: Black Hills [A]. Colorado: Steamboat Springs 

[USNM]. 

Pseudomicromus variolosus (Hagen) 

Micromus variolosus Hagen, Proc. Boston Soc. Nat. Hist., vol. 23, p. 284, 1886. 
California: San Bernardino Mts. [A]. Colorado: Cheyenne Can- 
yon [USNM]. Arizona: Chiricahua Mts. [A]; Tucson [USNM]. 

Pseudomicromus subanticus (Walker) 

Hemerobius subanticus Walker, Neuropt. British Mus., pt. 2, p. 282, 1853. 

California: Redding [USNM]. Arizona: Chiricahua Mts. [A]. 
Texas: BrownsviUe [USNM]. 

Genus Stenomicromus Kriiger 

Stenomicromus Kriiger, Stettin Ent. Zeit., vol. 83, p. 171, 1922. 

The dorsally spine-beset single aedeagus and the huge distal process 
of the anal plate are diagnostic of this genus. Parameres are separated, 
connected with each other only by a short bridge near base. The 
genus consists of the genotype, S. paganus of Europe, and its Nearctic 
counterpart, S. montanus. 

Stenomicromus montanus (Hagen) 

Micromus montanus Hagen, Proc. Boston Soc. Nat. Hist., vol. 23, p. 279, 1886. 
Alberta: Waterton Lakes National Park [A]. British Columbia: 
Alaska Highway [A]. Washington: Olympic National Park, Mt. 
Rainier, and Mt. St. Helena [A]. *Oregon: Wallowa Mts. and Willa- 
mette National Forest [A]. Colorado: Dream Lake and Glacier 
Creek in Rocky Mt. National Park, Gothic [A]. CaUfornia: Castle 
Crags [A]. 



212 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iia 

Genus Ameromicromus Nakahara 

Ameromicroynus Nakahara, Mushi, vol. 34, p. 33, 1960. 

This genus is endemic in North America, consisting of a single 
species, A. posticus. It is well characterized by the presence of a 
large subquadrate supraedeagal plate in the tenth sternite, which 
covers the aedeagus dorsally like a hood, and by the completely 
fused parameres, the apical two-thirds of which is in the form of a 
long pointed process and the basal one third, in that of a thin blade 
(Nakahara, 1960, p. 33). Kriiger's Paramicromus is a homonym. 

Ameromicromus posticus (Walker) 

Hemerobius posticus Walker, Neuropt. British Mus., pt. 2, p. 283, 1853. 

*Washington: St. Andrews Creek, 3800 ft., Mt. Ramier, July 17, 
1953, a single male [A]. 

Genus Hemerobius Linnaeus 

Hemerobius Linnaeus, Systema naturae, ed. 10, vol. 1, p. 549, 1758. 

All but two {H. nigrans and H. alpestris) of the previously recorded 
species of this genus are represented. There is an additional species 
in the USNM collection that I am unable to identify. 

Hemerobius humulinus Linnaeus 

Hemerobius humulinus Linnaeus, Systema naturae, ed. 10, vol. 1, p. 550, 1758. 

*Alaska: Teller [USNM]. *Washington: Mt. Hood and Alerritt 
[A]. This conmaon Holarctic species, previously recorded from British 
Columbia, seems to be very rare in western North America. 

Hemerobius pacificus Banks 

Hemerobius pacificus Banks, Trans. American Ent. Soc, vol. 24, p. 24, 1897. 

British Columbia: Wellington [USNM]. California: Kings Canyon 
National Park [A]; Mill Valley, Van Damme State Park, Inverness, 
Berkeley, Carmel, and San Jacinto Mts. [USNM]. Utah: Plam City 
and Logan [USNMJ. Arizona: White Alts, and Chiricahua Mts. [A]. 
New Mexico: Frijoles Canyon in Bandelier National Monument. 

In spite of the great difficulty in distinguishing the female of 
H. pacificus from that of H. neadelphns, I referred all the specimens 
to the former, which show accentuated asymmetry of the apical part 
of forewing with more pointed apex. 

Hemerobius neadelphus Gurney 

Hemerobius neadelphus Gurney, Ann. Ent. Soc. America, vol. 41, p. 214, 1948. 

British Colmnbia: Nanaimo and Wellington [USNM]. *Washing- 
ton: Mt. St. Helena, Olympic National Park, and Alt. Rainier [A]. 
Oregon: Blue Mts. [A]. California: Lost Creek in Lassen National 
Park, Hatchet Pass near Burne\", King's Canyon National Park, and 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 213 

Sequoia National Park [A]; Mill Valley [USNM]. *Montana: Gallatin 
City ]A]. *Colorado: Bierstadt Lake in Rocky Mountain National 
Park [A]. *Nevada: Washoe Co., and Kyle Canyon on Mt. Charleston 
[A]. *Arizona: Chiricahua Mts. [A]. 

There are over 50 females from various British Columbia, Alberta, 
California, Oregon, Utah, Wyoming, North Dakota, and Arizona 
localities that possibly may belong to this species but which I have 
not been able definitely to distinguish from H. pacifictos. As Gurney 
(1948) stated, there seems to be no dependable differential feature in 
the female between the two species. On the whole, the forewing 
seems to be slightly more rounded at apex in H. neadelphus than in 
H. pacificus. 

Hemerobius simulans Walker 

Hemerobius simulans Walker, Neuropt. British Mus.; pt. 2, p. 285, 1853. 

Alaska: Teklanika River in Mt. McKinley Park [A]. British 
Columbia: Alaska Highway and Kootenay National Park [A]; 
Wellington [USNM]. *Oregon: Rogue River National Forest, Lang- 
don Lake in Blue Mts., and Mt. Hood [A]. *California: Castle Lake 
[L^SNM]. *Montana: Avalanche Lake in Glacier National Park [A]. 
*Wyoming: Grand Teton National Park [A]. *Colorado: Clear Creek 
in Clear Creek Co. 

Hemerobius oralis Carpenter 

Hemerobius ovalis Carpenter, Proc. Amer. Acad. Arts Sci., vol. 74, p. 205, 1940. 
Alaska: Mt. McKinley National Park [A]. Washington: Mt. St. 
Helena [A]. Oregon: Rogue River National Forest [A]. California: 
Lassen National Park [A]. Wj^oming: Yellowstone National Park 
[A]. *Colorado: Bear Lake in Rocky Mountain National Park and 
Gothic [A]. 

Hemerobius stigmaterus Fitch 

Hemerobius stigmaterus Fitch, Noxious Ins. New York, reports 1 and 2, p. 93, 1856. 

Alberta : Waterton Lake National Park [A]. Washington : Cle Elmn 
[USNM]. Oregon: Bend, Lostine Valley in Wallowa Mts., Blue 
Mts., and Rogue River National Forest [A]. California: Yosemite 
National Park, Tioga Pass in Tuolumne Co., Nordon, and Lake 
Tahoe [A]; Fort Bragg, Miami Ranger Sta., Smith River, Castle Lake, 
San Jacinto Mts., Truckee, Keddie, and Nevada City [USNM]. 
*Montana: Glacier National Park [A]. Idaho: Twin Creek Camp in 
Salmon National Forest [A]. Wyoming: Teton Co. [USNM]. Colo- 
rado: Fraser [USNM]; Nymph Lake in Rocky Mountain National 
Park [A]. New Mexico: Chiricahua Mts. [USNM]. 

In a recent publication Tjeder (1960) synonymized H. stigmaterus 
under H. stigma Stephens, believing that the slight deviation in the 
shape of the median process of the tenth sternite (gonarcus) to be of 

734-073—64 2 



214 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

no specific value. He did not mention the difference in aedeagus, 
which to me seems rather important: H. stigma does not show the 
toothlike lateral expansion near the base that is present in H. stig- 
materus (see Gurney, 1948, fig. 10, and Nakahara, 1960, fig. 96). 
The intraspecific range of variability in these structm-es has not been 
explored adequately, however, and further studies involving dissection 
of a large number of specunens seems necessary in establishing this 
possible synonymy. 

Hemerobius conjunctus Fitch 

Hemerobius conjunctus Fitch, Noxious Ins. New York, reports 1 and 2, p. 94, 1856. 
Alaska: Mt. McKinley Park [A]. *Yukon: Alaska Highway [A]. 
Alberta: Banff National Park [A]; Mt. St. Piran [USNM]. *Wash- 
ington: Olympic National Park and Mt. St. Helena [A]. *Oregon: 
Blue Mountains, Crater Lake, and Wallowa Mts. [A]. *California: 
Yosemite National Park [A]. *Idaho: Twin Creek Camp in Salmon 
National Forest [A]. Utah: Uinta Mts. and Cedar Breaks National 
Monument [A]. *S. Dakota: Black Hills [A]. Colorado: Rocky 
Mountains National Park, Gothic, Pike National Forest, and Arapaho 
National Forest [A]. 

Hemerobius kokaneeanus Currie 

Hemerobius kokaneeanus Currie, Proc. Ent. Soc. Washington, vol. 6, p. 85, 1904. 
*Alaska: Prince of Wales Is. [USNM]. *Alberta: Waterton Lake 
National Park [A]. Washington: Olympic National Park and Mt. 
Rainier [A]. ^Oregon : Willamette National Forest and Mt. Hood [A]. 
♦Wyoming: Teton Co. [USNM]. Colorado: Newcastle [USNM]. 

Hemerobius bistrigatus Currie 

Hemerobius bistrigatus Currie, Proc. Ent. Soc. Washington, vol. 6, p. 79, 1904. 
British Columbia: Wellington [USNM]. *Washington: Mt. Rai- 
nier [A]; Baring [USNM]. Oregon: Ochoco Mts. [A]. Cahfornia: 
Lake Tahoe and Napa [USNM]. ^Montana: Glacier National 
Park [A]. 

Hemerobius species 

Alaska: Anchorage, one female [USNM]. 

Head, thorax, and abdominal tergites and sternites deep black, with 
a faint median light-colored streak on notum. Antennae dark brown; 
basal joint black. Forewing about 7 mm. in length; membrane uni- 
formly greyish, unmarked except for a dark dot on crossvein m-cu, 
and a dark longitudinal streak in pterostigmatic region. 

This may well be a new species. 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 215 

Hemerobius dorsatus Banks 

Hemerobius dorsatus Banks, Canadian Ent., vol. 36, p. 61, 1904. 

Alaska: Alaska Highway I A]. *Yukon: Alaska Highway [A]. Al- 
berta: Banff National Park and Jasper National Park [A];Kannanaskis 
[USNM]. British Columbia: Alaska Highway [A]. *Oregon: Crater 
Lake [A]. *Montana: Beaverhead National Forest [Aj. *Cahfornia: 
Yosemite National Park and Inyo National Forest [A]. Colorado: 
Rocky Mountain National Park [A]. 

Genus Brauerobius Kriiger 

Brauerobius Kriiger, Stettin Ent. Zeit., vol. 83, p. 171, 1922. 

This genus is characterized by the exceedingly elongated anal 
plate of the male, which is rounded apically and bears no spinous pro- 
jection (Nakahara, 1960, p. 50). Internal ventral margin of the 
plate is beset with numerous denticulate tubercles toward apex. It 
consists of three species: G. marginatus (Stephens), type species, G. 
tristriatus (Kuwayama), and G. costalis (Carpenter), the last being 
the sole Nearctic representative. 

Brauerobius costalis (Carpenter) 

Hemerobius costalis Carpenter, Proc. Amer. Acad. Arts Sci., vol. 74, p. 213, 1940. 

Alaska: Fairbanks [USNM]. British Columbia: Yoho, Banff, and 

Waterton Lakes National Parks [A]. *Idaho: Boise National Forest 

[A]. 

Genus Kimminsia Killington 

Kimminsia Killington, Monogr. British Neuropt., vol. 2, p. 254, 1937. 

Eleven species of this genus are recognized in the material before me, 
and three of them are described as new. These three, though repre- 
sented by a single specimen each, one unfortunately a female, are of 
such distinctive characters that I feel entirely safe in naming them. 
These were all collected by Dr. Alexander. There are three other 
species of this genus previously recorded from western North America 
but not found in the material : K. Jumata Carpenter, K. longipennis 
(Banks), and K. constricta Parfin. 

Kimminsia disjuncta (Banks) 

Hemerobius disjuncta Banks, Trans. Amer. Ent. Soc, vol. 24, p. 25, 1897. 
Alaska: Matanuska [USNM]. British Columbia: Kaslo [USNM]. 

Kimminsia coloradensis (Banks) 

Hemerobius coloradensis Banks, Trans. Amer. Ent. Soc, vol. 24, p. 26, 1897. 
Washington: Blue Mts. [USNM]. 



216 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Kimminsia involnta Carpenter 

Kimminsia involuta Carpenter, Proc. Amer. Acad. Arts Sci., vol. 74, p. 219, 1940. 
Alaska: Fairbanks [USNM]. British Columbia: Kokanee Mts. 
[USNM]. 

Kimminsia brunnea (Banks) 

Boriomyia brunnea Banks, Bull. Mus. Comp. Zool. Harvard, vol. 64, p. 333, 1920. 
Alaska: Mt. McKinley Park [A]. Alberta: Jasper and Banff Na- 
tional Parks [A]. Colorado: Head of Elk Creek [USNM]. 

Kimminsia pretiosa (Banks) 

Boriomyia praetiosa Banks, Trans. Amer. Ent. Soc, vol. 34, p. 260, 1908. 

Colorado: Clear Creek [USNM]. Utali: Uinta Canyon and Lo- 
gan [USNM]. 

Kimminsia schwarzi (Banks) 

He{merobius schwarzi Banks, Proc. Ent. Soc. Washington, vol. 5, p. 241, 1903. 

*Alaska: Mt. McKinley Park [A]. California: Yosemite National 
Park [A]. Arizona: Williams [USNM]. 

Kimminsia olympica, new species 

Figure 3; Plate 1 (Figure 3) 
Holotype d', Deer Park (5400 ft.), Olympic National Park, Wash- 
ington, July 17, 1948 (C. P. Alexander). Right forewing (dry) and 







Figure 3. — Kimminsia olympica new species; a, terminal abdominal segments, lateral 
view; b, the same, dorsal view; c, apical comb of anal plate, internal view; d, tenth 
sternite, lateral view; e, parameres, lateral view. 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 217 

dissected parts of genitalia (in balsam) mounted on two slides. 
USNM type 66176. 

Face uniformly fuscous brown; vertex with two elongated pale 
patches divided by median fuscous brown longitudinal stripe; palpi 
and antennae brownish. Pronotum broadly brownish yellow medi- 
ally, darker on sides. Mesonotum brownish yellow, fuscous brown 
on sides. Metanotum almost totally fuscous brown. Abdomen 
fuscous brown. Legs brownish yellow. 

Forewing: Length 9.5 mm, width 3.5 mm, rather elongated with 
rounded apex; longitudinal veins pale with fuscous brown spots and 
short streaks; inner and outer gradates deeply fuscous and strongly 
margined with brown; basal crossveins m-cu and cu also fuscous and 
margined; membrane hyahne, with many small sagittate maculations 
in discal area; outer and hindmarginal area marked with scattered 
brownish patches; the markings forming a long brown fascia across 
the wing over inner gradates, one over the basal crossveins, a short 
one in hindmarginal area between the two, and a fom*th, interrupted 
fascia over outer gradates. Hindwing hyaline; veins fuscous, except 
toward base, where they are pale. 

Male genitalia: Anal plate long and slender, rounded apically, with 
a stout sharply pointed ventroapical process, which is bent strongly 
forward and somewhat inward. Tenth sternite rather narrow, dorsal 
bridge between "wings" short, narrowly produced posteriorly, bearing 
long and laterally compressed aedeagus over it; ventral process very 
long and slender. Parameres turned up at both ends, fused in middle; 
the^separated basal parts short, and distal parts very much longer. 

This species is like Kimminsia fumata, K. comtricta, K. pretiosa, 
and K. schwarzi in having down-curved ventroapical process to anal 
plate, but the anal plate itself is much longer and more slender in this 
species, and the process is strongly bent forward and is sharply pointed. 
It is a rather conspicuous species with maculations roughly formmg 
four transverse bands across the forewing. 

Kimminsia posticata (Banks) 

Boriomyia posticata Banks, Trans. Amer. Ent. Soc, vol. 32, p. 39, 1905. 

*Yukon: Alaska Highway [A]. ^California: Tuolumne Meadows 
m Yosemite National Park [A]. Utah: Logan [USNM]. 

Kimminsia alexanderi, new species 

Figure 4; Plate 1 (Figure 4) 

Holotype cf, Haines Highway, Alaska, July 5, 1952 (C. P. Alex- 
ander). Right forewing (dry) and dissected parts of genitalia (in 
balsam) mounted on two slides. USNM type 66177. 

Face fuscous brown, with a narrow yellow transverse line along the 



218 



PROCEEDINGS OF THE NATIONAL MUSEUM 



base of clypeus, vertex yellow; palpi brown; antennae brow^nish 
yellow, darker beyond middle. Pronotum brownish yellow, mth 
fuscous-brown longitudinal stripe in middle and on sides; meso- and 
metanotum broadly yellowish medially, browTiish on sides. Legs 
yellowish. Abdomen yellowish basally, brownish beyond middle. 




Figure 4. — Kimminsia alexanderi, new species: a, terminal abdominal segments, lateral 
view; b, the same, dorsal views; c, apical comb of anal plate, internal view; d, tenth 
sternite, lateral view; e, parameres, lateral view. 



Forewing: Length 7 mm., width 3 mm.; apex rounded; membrane 
very slightly smoky, practically immaculate in discal area; outer 
marginal area down to the end of cubitus with alternate brownish 
and hyaline small patches; the patches larger in hindmarginal area 
from Cui to hindmargin. A rather conspicuous dark-brown spot 
over the last crossvein of the inner gradate series. Longitudinal 
veins fuscous, interrupted with short pale streaks; crossveins of the 
same shade of fuscous. Hindwing sUghtly smoky with fuscous vena- 
tion, which is paler toward base. 

Male genitalia: Anal plate very long and narrow, distally curved 
roundly downward and then forward; apical part provided with a 
closely set series of short teeth on ventrolateral margin. Tenth 
sternite produced posteriorly into a large subtriangular lobe (in 
dorsal view), ending in a short pointed dorsoapical process; lateral 
"wings" broad, with a large lateroposterior process, which ends in 
inwardly curved obtuse apex. Parameres slender, strongly curved, 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 219 

fused in middle and separated proxiinally and distally; the proximal 
lobes shorter than distal parts, all sharply pointed at ends. 

The strongly curved ventroapical process of anal plate in this 
species reminds one of K. posticata, but anal plate is very much 
narrower in lateral view and the apical prolongation is only slightly 
more slender than the body of the plate. This species is very much 
smaller than K. posticata, and the forewing membrane is uniformly 
smoky in discal area, maculations being confined to postcubital area. 

Kimminsia furcata (Banks) 

Boriomyia furcata Banks, Psyche, vol. 42, p. 55, 1935. 

Alaska: Toklat Kiver [USNM], Haines Highway, Mt. McKinley 
National Park, and Richardson Highway [A]. Alberta: Jasper 
National Park and Kicking Horse Pass [A]. *British Columbia: 
Alaska Highway [A]. * Washington: Olympic National Park [A]. 
California: Yosemite National Park, Tioga Pass in Tuolumne Co, [A]. 
*Nevada: Lake Tahoe [A]. Colorado: Timber Creek Camp, Cham- 
bers Lake, and Monarch Pass in Rocky Mt. National Park. [A]. 

Kimminsia melaleuca, new species 

Plate 1 (Figure 5) 

Holotype 9, Alaska Highway, Mile Post 720, Yukon Territory, 
July 2, 1952 (C. P. Alexander). Right forewing mounted dry on a 
slide. USNM type 66178. 

Face shining black, brownish toward anterior margin of clypeus; 
vertex dull black. Antennae fuscous, almost black. Pronotum 
black, narrowly margined with brownish yellow anteriorly, with two 
ill-defined brownish-yellow longitudinal stripes submedially. Meso- 
notum black, with two brownish-yellow patches lateroposteriorly, 
each enclosing a short black longitudinal streak. Metanotum and 
abdomen black. Legs brownish yellow, femur broadly banded with 
dark brown toward both ends. 

Forewing: Length 7 mm., width 3 mm., apex rounded. Veins 
black, interrupted with numerous white spaces; both series of gra- 
dates deep black, strongly margined with fuscous black, especially the 
posterior crossveins of the inner series ; sagittate maculations of discal 
area strongly developed and mostly transversely confluent; outer and 
hindmarginal areas, distal to outer gradates, and behind Cui decorated 
Avith large patches of alternate black and hyaline white. Maculations 
faint in costal space. Hind wing very distinctly smoky, with fuscous 
black venation; crossveins in pterostigmatic area conspicuously 
white. 

The forewing of this black-bodied species is very striking because of 
the hyaline spots that appear so white owing to the whiteness of the 



220 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii6 

parts of the veins enclosed, in strong contrast to the deeply fuscous- 
black markings. 

An attempt has been made to accommodate the three new species 
described above in Carpenter's key (1940, p. 215) and its partial 
modification by Parfin (1956, pp. 207-208). 

Key to the Nearctic Species of Kimminsia 

1. Pronotum with a conspicuous median yellowish stripe, bordered laterally 

with dark brown 2 

Pronotum without such a median stripe 10 

2. Upper part of frons very dark brown, lower part yellowish or light bro\\Ti 

with very abrupt transition K. coloradensis Banks 

Frons more uniformly dark brown, or, if the upper part is darker than the 
lower, the transition is very gradual 3 

3. Forewing with blackish-brown spots at distal m-cu crossvein and from Cui 

to hindmargin 4 

Forewing with maculations more evenly distributed 5 

4. Forewing nearly uniformly smoky in discal area; small species (forewing 

7 mm.) K. alexandcri, new species 

Forewing with some sagittate maculations in discal area; larger species 
(forewing 10 mm.) K. posticata Banks 

5. Anal plate of male with a very long dorsal process. . . . K. furcata Banks 
Anal plate of male without dorsal process 6 

6. Anal plate of male with very long and slender apical process that is terminally 

expanded K. subnebulosa (Stephens) 

Anal plate of male with short apical process 7 

7. Process of anal plate strongly bent forward . . . K. olympica, new species 
Process of anal plate directed more or less straight downward 8 

8. Process of anal plate short and bent inward K. constricta Parfin 

Process of anal plate moderately long, not bent inward 9 

9. Large species (forewing averaging 11 mm.), costal area broad. 

K. involuta Carpenter 

Smaller species (forewing averaging 9 mm.), costal area of moderate 

breadth K. disjuncta (Banks) 

10. Face, vertex, and pronotum deep black . . . . K. melaleuca, new species 
Face, vertex, and pronotum yellowish or light brown 11 

11. Face, vertex, and pronotum with scattered red specks . K. longipennis Banks 
Face, vertex, and pronotum not so marked 12 

12. Forewing without maculation K. brunnea Banks 

Forewing with conspicuous maculations 13 

13. Forewing without distinct transverse bands K. pretiosa Banks 

Forewing with distinct transverse bands 14 

14. Mesonotum with dark-brown anterior border K. schwarzi Banks 

Mesonotum uniformly light brown K. fumata Carpenter 

Genus Wesmaelius Kriiger 

Wesmaelius Kruger, Stettin Ent. Zeit., vol. 82, p. 170, 1922. 
There is a single Nearctic species of this genus. 



HEMEROBIIDAE OF WESTERN NORTH AMERICA — NAKAHARA 221 

Wesmaeliiis longifrons (Walker) 

Hemerobius longifrons Walker, Neuropt. British Mus., pt. 2, p. 291, 1853. 

Oregon: Princess Creek Forest Camp on Odell Lake in Klamath 
Co. [A]. *Idaho: Twin Creek Camp in Salmon National Forest and 
Lolo National Forest [A]. *Montana: Glacier National Park [A]. 
Colorado: Cheyenne Mts. [USNM]. 

Genus Megalomus Rambur 

Megalomus Rambur, Hist. Nat. Ins., Neuropt., p. 418, 1842. 
Only one of the four Nearctic species of this genus is represented. 

Megalomus moestus Banks 

Megalomus moestus Banks, Trans. Amer. Ent. Soc, vol. 22, p. 314, 1895. 
Arizona: Chiricahua Mts. [A and USNM]. 



Literature Cited 

Banks, Nathan 

1895. New Neuropteroid insects. Trans. Amer. Ent. Soc, vol. 22, 

pp. 313-316. 
1897. New North American Neuropteroid insects. Trans. Amer. Ent. Soc, 

vol. 24, pp. 21-31. 

1903. Neuropteroid insects from Arizona. Proc. Ent. Soc. Washington, 

vol. 5, pp. 237-245. 

1904. A list of Neuropteroid insects, exclusive of Odonata, from the vicinity 

of Washington, D.C. Proc. Ent. Soc. Washington, vol. 6, pp. 201- 
217. 
1904. New species of Hemerobius. Canadian Ent., vol. 36, pp. 61-63. 

1904, Neuropteroid insects from New Mexico. Trans. Amer. Ent. Soc, 

vol. 30, pp. 97-110. 

1905. A revision of the Nearctic Hemerobiidae. Trans. Amer. Ent. Soc, 

vol. 32, pp. 21-51. 
1908. Neuropteroid insects: Notes and descriptions. Trans. Amer. Ent. 

Soc, vol. 34, pp. 255-267. 
1911. Descriptions of new species of North American Neuropteroid insects. 

Trans. Amer. Ent. Soc, vol. 37, pp. 335-360. 
1920. New Neuropteroid insects. Bull. Mus. Comp. Zool., Harvard Coll., 

vol. 64, pp. 299-362. 
1935. A few new North American Neuroptera. Psyche, vol. 42, pp. 53-57. 

Carpenter, F. M. 

1940. A revision of the Nearctic Hemerobiidae, Berothidae, Sisyridae, 
Polystoechotidae, and Dilaridae (Neuroptera). Proc Amer. Acad. 
Arts Sci., vol. 74, pp. 193-280. 



222 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii6 

CURRIE, R. p. 

1904. Notes on some Hemerobiidae from Arizona and California. Proc. 

Ent. Soc. Washington, vol. 6, pp. 79-81. 
1904. Hemerobiidae from the Kootenay district of British Columbia. Proc. 

Ent. Soc. Washington, vol. 6, pp. 81-90. 

Fitch, Asa 

1856. First and second report on the noxious, beneficial and other insects 
of the state of New York. Albany, 336 pp. 

GuRNEY, Ashley B. 

1948. Notes on Nearctic Hemerobiidae, with descriptions of two new species 
(Neuroptera). Ann. Ent. Soc. Amer., vol. 41, pp. 213-222. 

Hagbn, Herman A. 

1886. Monograph of the Hemerobiidae. Proc. Boston Soc. Nat. Hist., 

vol. 23, pp. 250-292. 

KiLLINGTON, F. J. 

1937. A monograph of the British Neuroptera, vol. 2, 306 pp. 

KRtJGER, L. 

1922. Beitrage zu einer Monographie der Neuropteren — Familie der 
Hemerobiiden. Stettin Ent. Zeit., vol. 83, pp. 138-172. 

Linnaeus, C. 

1758. Systema naturae, ed. 10, vol. 1. 

McLachlan, R. 

1869. New species, etc., of Hemerobiina ; with synonymic notes (first 
series). Ent. Monthly Mag., vol. 6, pp. 21-27. 

Nakahara, Ward 

1960. Systematic studies on the Hemerobiidae. Mushi, vol. 34, pp. 1-69. 

Parfin, Sophy 

1956. Taxonomic notes on Kimminsia (Neuroptera: Hemerobiidae). Proc. 
Ent. Soc. Washington, vol. 58, pp. 203-209. 

Rambur, p. 

1842. Histoire naturelle des insectes: Nevroptferes. 

Stephens, J. F. 

1836. Illustrations of British entomology. Mandibulata, vol. 6. 

Tjeder, B. 

1960. Neuroptera from Newfoundland, Miquelon, and Labrador. Opuscula 
Ent., vol. 25, pp. 146-149. 

Walker, F. 

1853. List of the specimens of Neuropterous insects in the collection of the 
British Museum, pt. 2. 



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FoREWTXGS. — 1, Sympkerobius brunneus, new species; 2, Sympherobius lexanus, new species ; 
3, Kimminsia olympica, new species; 4, Kimminsia alexanderi, new species; 5, Kimminsia 
melaleuca, new species. (Relative sizes of wings not shown.) 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1965 Number 3504 

A CONTRIBUTION TO THE STUDY OF 

THE GENUS SPHAEROCERA LATREILLE 

IN CENTRAL AND SOUTH AMERICA 

(DIPTERA: SPHAEROCERIDAE) 



By O. W. Richards ' 



The latest revision of the American flies of the genus Sphaerocera 
Latreille (Malloch, 1925) records only one species of this genus from 
south of Panama: S. flavicoxa Malloch from Brazil and Costa Rica. 
Malloch records four species (S. flaviceps Malloch, S. pallipes Malloch, 
S. bimaculata Williston, and S. varipes Malloch) from Central America 
and the West Indies. Spuler erected a subgenus Parasphaerocera for 
S. bimaculata WiUiston that Malloch did not accept because it was 
founded principaUy on a color character to which two species, S. 
annulicornis Malloch and S. pallipes Malloch, though allied to the 
others, were an exception. In a re-examination of the genus, how- 
ever, I find that there are four distinct groups, each with four or more 
species, and it is probably convenient to treat each group as a subgenus. 

Key to Subgenera of Sphaerocera Latreille 

1. Scutellum with, at most, one basal marginal tooth on each side. Mesoscutum 
with rows of minute bristles. Abdominal sternites nearly always more or 
less reduced, especially in female. Hindtibia with or without apical 
spur 2 



' Department of Zoology and Applied Entomology, Imperial College, Univer- 
sity of London, England. 

223 



224 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Scutellum with six or more denticles along posterior margin. Mesoscutum 
with rows of warts. Abdominal sternites broad. Hindtibia with apico- 
ventral spur. Face with central elevation that forms a sharp lower 
boundary to the antennal foveae 3 

2. Face not elevated below, antennal foveae not sharply defined. Abdomen 

without dorsal pale spots .... Su'ogenus Sphaeroccra Latreille 1804, 

Type by subsequent monotj'py, Sphacrocera curvipes Latreille, 1805. 

(Other species: S. monilis Haliday, 1836; <S. flaviceps Malloch, 1925; 

S. jeanneli Richards 1938; S. uittei Vanschuytbroeck, 1948; S. ruandana 

Vanschuytbroeck, 1948; S. longipes Richards, 1951.) 
Face elevated so that the antennal foveae are well defined below by a sharp 
keel. Abdomen usually with two large whitish spots. 

Subgenus Parasphaerocera Spuler, 1924 

Type by original designation, Sphaerocera himacidata WiUiston, 1896. 

(Other species: S. annulicornis Malloch, 1913; (S. pallipes Malloch, 1914; 

S. flavicoxa Malloch, 1925; S. nigrifemur Malloch, 1925; S. varipes 

Malloch, 1925; S. striata Malloch, 1925; S. galapagensis Curran, 1934; 

and the new species described below.) 

3. Vein M1+2 and, to a less extent, R4+5 strongly bent forward. Mesoscutum 

uniformly covered with warts, longitudinal bare lines indistinct or absent. 

Subgenus Lotobia Lioy, 1864 
Type by monotypy, Borborus pallidiventris Meigen, 1830. (Other 
species: Sphaerocera simia Seguy, 1933; S. arcuata Seguy, 1933; S. 
kivuenis Vanschuytbroeck, 1948; S. ruishuruensis Vanschuytbroeck, 
1948; and seven species described by Vanschuytbroeck in 1959.) 
Veins M1+2 and R4+5 not bent forward. Mesoscutum with rows of warts and 
wide bare spaces in between some of the rows. 

Subgenus Ischiolepla Lioy, 1864 
Type by monotypy, Borborus denticulalus Meigen, 1S2>Q = Sphaerocera 
nitida Duda, 1923. (Other species: Sphaerocera pusilla (Fallen, 1820); 
*S. scabricula Haliday, 1836; S. vaporariorum Haliday, 1836 = 5. para- 
pusilla Duda, 1923; S. orientalis de Meijere, 1908; S. scabra Spuler, 
1924; S. jansseni Vanschuytbroeck, 1948; S. flava Vanschuj'tbroeck, 
1951; S. dura Vanschuytbroeck, 1959; S. kifaruensis Vanschuytbroeck, 
1959; S. crenata (Meigen, 1838) =<S. coronata (Zetterstedt, 1838); S. 
paracrenata Duda, 1923; S. falcozi Duda, 1921; <S. micropyga Duda, 
1923. The last four species form a separate subgroup.) 

Subgenus Parasphaerocera Spuler, 1924 

This subgenus appears to be exclusively American and it now appears 
that the species, though superficially similar to one another, are really 
very numerous. When I was in Berkeley in 1961, I was able to 
study a long series of males and females from one locality in Ecuador, 
a series that was preserved in the collection of the Cahfornia Academy 
of Sciences (CAS) . At first, the series seemed to consist of one species, 
but more careful study showed that there were three. In order to 
discover the relationship of these species with the supposedly common 
Central American species, S. bimaculata Williston, I borrowed all 
the specimens resembling that species in the collection of the U.S. 
National Museum (USNM). Through the kind help of Dr. J. F. 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 225 

Gates Clarke and Mr. Curtis Sabrosky, 27 specimens were sent to 
me. Careful study showed that 9 species were represented among a 
random distribution of males and females. MaUoch first pointed 
out that the reduced sternites of this group provide specific characters. 
It seems that they provide by far the most important characters and 
require careful study in every form. Unfortunately, the type of 
reduction seems to be quite different in the two sexes, which are very 
difficult to associate. In some species the structure of the pro- 
sternum is a guide. The presence or absence of a minute antero ventral 
spur on the hindtibia is less useful than MaUoch supposed because it 
may vary in size and, in some species, is difficult to detect. The 
nomenclature of the parts of the genitalia is the same as in Richards 
(1961). The new species described below are compared with S. 
ecuadoria, new species, rather than with S. bimaculata Williston 
because only in the former is a long series of both sexes available. 
Single specimens were examined of what appear to be six other species, 
but their characters are largely minor deviations from those of the 
species that are described. 

New Species AUied to Sphaerocera bimaculata Williston 
Sphaerocera {Parasphaerocera) ecuadoria, new species 

Males and females. Black, dull, mesoscutum and pleuron only 
slightly more shining; antennae brown; mentum and palpi testaceous; 
legs pale yellow brown, tarsi and apices of femora slightly darker. 
Abdomen with two large pale spots. Halteres yellow. Wings 
h^^^line, venation brown. Length 2.5-3.0 mm. 

Lower face forming a flat trapeziform plate (fig. 7), coming up to 
a point between the antennae; area of lunula also raised into a flat 
triangular plate whose point meets that of the lower face at about the 
lowest point of the antennal attachment; antennal foveae thus sharply 
defined by a keel that is especially high internally and ventrally. 
Buccae not striate except rather indistinctly posteriorly. Arista 
four times as long as antenna with very short pubescence. Dorsum 
of head very dull, no proper bristles. Prosternum (fig. 6) in the form 
of a linear projection, about half as long as width of forecoxae, from 
the triangular depression in front of the mesosternum. Mesoscutum 
with some scaly tomentum and four indistinct rows of very short 
bristles (corresponding approximately to dorsocentrals and acrosti- 
chals), a few further, more lateral, bristles, irregularly placed. Scu- 
tellum with a smaU denticle on each side, where the hindmargin curves 
around to become transverse, doreal surface almost bare. Legs 
thick, especially in the c^, d^ hindfemur twice as thick as midfemur. 
Hindtibia with a small black anterior apical spur, shorter than one- 



226 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

quarter the apical width of tibia, spur smaller and paler in the female. 
Hindbasitarsus longer than the second segment but not as long as 
2 + 3. Abdomen with tergal plates 1+2, 3, and 4, each slightly in- 
creasing in size; pale spots (fig. 1) caused by desclerotization of cuticle, 
lying across the boundaries between 1 + 2 = 3, and 3=4; anterior spot 
oval, posterior one pointed laterally, both well separated from the 
margin. Female with segments after 4 (figs. 2, 3) often retracted but 
tergite 5 sometimes visible as a narrow transverse plate; sternites 
(fig. 3) very reduced, 1+2 rather small and transverse, 3 somewhat 
longer than broad, weakly dumbbell shaped (or in the Chilcales 
specimen, diamond shaped), 4 very long and narrow, more than four 
times as long as broad, side margins sinuate; sternite 5 transversely 
crescentic; spu-acles 1-4 in the membrane laterally (1 hard to see 
and very close to the thorax), 5 in the membrane between the fifth 
sternite^ and tergite; cerci oval with short bristles. Male with ter- 
gite 5 relatively large, somewhat better developed on the right, ter- 
gites 6 and 7 visible on the left side; genitalia (figs. 4, 5) large; ninth 
segment not emarginate for the reception of the cerci and the anal 
orifice; cerci loosely attached to the ninth segment, emargination 
between them shallow, flat bottomed; gonapophyses absent, lateral 
lobes of forked plate short and broad, lying at the sides of the cerci; 
forceps long, narrow, pointed, curved upward, yellowish with rather 
long bristles on the inner sides; aedeagus with a basal stalk that ex- 
pands distally and is angularly truncate, distally with an apical loop- 
like sclerite, each side of which is a lateral lobe; sternites 1+2 (fig. 4) 
represented by three small plates, the central one transverse and big- 
ger than the other two; sternite 3 transveree with a moderately long 
and broad rounded posterior extension; sternite 4 very transverse, 
narrowed laterally; sternite 5 even more transverse with a central 
desclerotized area; sternites 3 and 4 with some longish bristles. 

Ecuador: Los Kios, Pichilingue, elev. 40 m., Feb. 2, 1955, 20 cT 
18 9 (including holotype cf and 9 allotype), E. S. Ross and E. I. 
Schlinger, CAS; further paratypes, Guayas, Naranjal, December 
1955, 9, Chilcales, Aug. 1, 1955, 9, R. Levi-Castillo, USNM. 

Some paratypes in my collection, including Icf , 1 9, macerated and 
in alcohol. The females and males are associated because they were 
the common type in the locality with the same type of prosternum. 

Sphaerocera {Parasphaerocera} trapezina, new species 

Males and females. Resembles S. ecuadoria, new species, very 
closely but differs as follows: Size on the average slightly smaller. 



2 Young (1921, p. 260) found seven spiracles in Copromyza ( = Borborus) 
equina Fall. I could find only five in C. sordida Zetterstedt, c?" . 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 227 




Figures 1-7. — Sphaerocera ecuadoria, new species: 1, Dorsal view, cf abdomen. 2, Dorsal 
view, 9 abdomen, tergite 5 to end extended. 3, Ventral view, 9 abdomen, distal part 
extended, spermathecae drawn at sides. 4, Ventral view, cf abdomen: a, forceps; b, 
cerci; c, lateral lobe of forked plate. 5, Genitalia, c?, from the right: a, forceps; b, cerci; 
c, lateral lobe of forked plate. 6, Prosternum, c?". 7, Facial plate, cf • 



228 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iis 

Anterior abdominal spot somewhat smaller, separated by half its own 
width from the margin. Prosternum slightly wider, especially pos- 
teriorly, and somewhat shorter, Hindtibia with anterior apical spm* 
very minute, almost obsolete. Male genitalia (figs. 8, 9) large, 
ninth segment feebly emarginate to receive the cerci. Cerci loosely 
articulated to the ninth segment with a deep U-shaped emargination 
between them. Lateral lobes of forked plate narrower and more 
pointed at the apex. Forceps long, narrow, pointed, curved down- 
ward, pale yellow with denser, shorter bristles. Aedeagus generall}' 
similar but basal stalk blunter, apical part produced into a strong 
proximal angle, lateral lobe less developed. Sternites 1+2 (fig. 8) 
represented by three small plates, the central one very small, smaller 
than the other two (sometimes rather longer and more transverse 
than in fig. 8); sternite 3 trapeziform, posterior extension shorter and 
wider than in S. ecuadoria; sternite 4 very transverse, not narrowed 
laterally, sides nearly straight; sternite 5 rather irregular with a 
considerable central membranous emargination; sternite 3 almost 
bare, sternite 4 with some long bristles. Female (fig. 10) with sternites 
1 -}-2 often represented by a very small transverse sclerite, sometimes 
absent; sternite 3 usually absent, sometimes represented by a sclero- 
tized dot; sternite 4 about two and one-half times as long as its 
greatest width, drop shaped, narrower anteriorly than posteriorly, 
where it is rounded, exact shape a Httle variable; sternite 5 transversely 
crescentic, about as in S. ecuadoria. 

Ecuador: Los Kios, Pichilingue, elev. 40 m., Feb. 2, 1955, 5cf, 8 9 
(including holotype c? and allotype 9), E. S. Ross and E. I. Schlinger, 
CAS. Paratypes: Ecuador: Los Rios 27 mi. SW. Quevedo, elev. 
50 m., Feb. 5, 1955, cf , Ross and Schlinger, CAS; Guayas, Naranjal, 
December 1955, cf, R. Le\^-CastiUo, USNM. Some paratypes, 
including one macerated cf , are in my collection. 

Sphaerocera (Parasphaerocera) tertiOf new species 

Male. Resembles S. ecuadoria, new species, very closely but differs 
as foUows: Length about 2.6 mm. Prosternum slightly wider with 
indications of a transverse basal piece from which the linear pro- 
jection arises. Abdominal spots broader and the posterior one also 
longer, about as S. trapezina. Genitalia (figs. 11, 12) large, ninth 
segment feebly emarginate for the reception of the cerci; cerci loosely 
attached to the ninth segment, emargination between them deep 
and almost V-shaped; basal lobes of forked plate longer and more 
pointed than in S. ecuadoria; forceps long, narrow, pointed, yellow, 
curved downward more strongly than in aS". trapezina, with numerous 
setae; aedeagus with a short basal stalk bearing finger-shaped proc- 
esses on each side, distally with a double lateral lobe on each side and 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 229 

a looplike apical piece; sternites 1+2 (fig, 11) represented by three 
small plates, aU about the same size, the central one usually very 
transverse; sternite 3 angularly dumbbell shaped but narrower 
posteriorly than anteriorly, the exact size and shape varying very 
little; sternite 4 very transverse, approximately rectangular, with a 
few short bristles; sternite 5 irregular, with a central membranous 
emargination. 

Ecuador: Los Rios, Pichilingue, elev. 44 m., Feb. 2, 1955, holo- 
tyi)8 and para type, cf , Ross and Schlinger, CAS. Paratypes: Panama 
CanrJ Zone: Smnmit, September 1946, cf , N. H. L. Krauss, USNM. 
Costa Rica: San Mateo, Higuito, cT macerated, Pablo Schild, USNM. 
Mexico: Vera Cruz, Cordoba, Apr. 13, 1908, &, Dr. A. Feynes, 
USNM. 

One of the Los Rios specimens is in my collection. 

Spliaerocera iParasphaerocera) chimborazo, new species 

Males and females. Generally like S. ecuadoria, new species, 
but differs as follows: Legs slightly darker though still yellow brown. 
Size slightly larger, length at least 3.0 mm. Facial plate raised into 
a convex rounded boss, quite conspicuous in profile. Prosternum 
consisting of a small posterior triangle, of which the tip is rounded 
in the female, linear projection scarcely sclerotized. Legs somewhat 
longer and less thickened, even in the cf, hindtibial spm- scarcely 
detectable; cf forefemm* with a row of short outstanding ventral 
bristles on proximal half. Abdominal spots somewhat larger. Male 
genitalia rather large, with tergite in profile somewhat angularly 
produced, evidently somewhat angularly emarginate, but most of the 
structures hidden; sternites 1+2 (fig. 13) represented by a single 
pea,r-shaped central spot; sternite 3 narrowly rectangular, very 
slightly widened at each end ; sternite 4 generally rectangular, slightly 
widened posteriorly, about twice as long as broad, produced spout- 
like beneath the genitalia; sternite 5 hidden. Female with anterior 
abdominal spot more produced at sides posteriorly than in male; 
sternite 1+2 (fig. 14) as in cT, sternite 3 not sclerotized, sternite 4 
more than twice as long as broad, somewhat narrowed at each end, 
sternite 5 trapeziform, anterior and posterior margins straight, sides 
somewhat concave; cerci oval, yellowish, with a moderate dorsal 
and apical bristle. 

Ecuador: Chimborazo, Bugna, holotype cT, allotype 9, R. Levi- 
Castnio, USNM type no. 66592. 

The sexes easily can be associated in this species by the structure 
of the face and prosternum. 



230 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Sphaerocera iParasphaerocera) bimaculata Williston 

Sphaerocera bimaculata Williston, 189G, Trans. Ent. Soc. London, p. 435. 

This species was described from St. Vincent; two females, both 
marked as co-types, are in the British Museum collection (BM), 
together with another female from Grenada. I have no reliable 
evidence at present that the species occurs elsewhere; all older 
records from Florida and Central America require confirmation. 

Female. Close to S. ecuodoria, new species, but differing as follows: 
Legs uniformly bright yellow brown. Size slightly smaller. Ab- 
dominal pale spots larger, both wider and longer. Prosternum \vith 
a narrow very transverse posterior piece, almost fused with the 
mesosternum, and a short almost linear anterior process, which in one 
9 is hardly sclerotized. Hindtibial spur very small indeed. Hind- 
basitarsus a fraction larger. Sternites 1+2 (fig. 15) represented by a 
minute sclerotized point; sternite 3 small, subrectangular, three times 
as long as broad; sternite 5 subcrescentic; cerci yellow, each with 
about 6 rather long hairs. 

Windward Islands: St. Vincent, 2 9 co-types, H. H. Smith; Grenada, 
Balthazar, 9, H. H. Smith, BM. 

Among the S. bimaculata are IcT and 19 (latter headless) also from 
Balthazar but apparently belonging to Si^haerocera (s.s.) species. The 
abdomen has no pale spots; the face is partly orange, the antennal 
foveae rather deep, with a distinct prominence below, on which stands 
a short but rather stout bristle; ventral hindtibial spur long but weak 
and rather hairlike. A female close to S. bimaculafa but apparently 
different comes from Panama Canal Zone (Barro Colorado Island, 
July 16, 1923, R. C. Shannon). In it the sides of sternites 3 and 4 
are straight, not sinuous, and sternite 4 is considerably longer. 

Sphaerocera {Parasphaerocera) levicastilli, new species 

Alale. Resembles S. ecuadoria, new species, very closely but differs 
as follows: Coxae and femora whitish, apical third of femora, tibiae, 
and tarsi light brown. Prosternum (fig. 17) consisting of a small 
equitriangular plate attached to the mesosternum with a narrow 
projection forward nearly twice as long as plate. Legs not thickened, 
hindtibiae with no anteroventral spur. Male genitalia large, with 
tergite rather distinctly emarginate to receive the cerci, which are 
separated by a deep V-shaped emargination; details of genitaha con- 
cealed. Sternites 1+2 (fig. 16) consisting of two round sclerotized 
dots and a small triangular plate; sternite 3 sub triangular, narrowing 
posteriorly but posterior end rounded, length less than twice the 
proximal width; sternite 4 sub triangular, quite wide, less than twice 
as long as wide, posterior end slightly narrowed before apex and 
weakly emarginate; sternite 5 almost completely hidden, transverse. 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 231 

O O O O 

o 

17 




10 



^ 






o 







14 



15 



n 



z:^ 



17 



16 



Figures 8-17. — Sphaerocera trafezina, new species: 8, Abdominal sternites and genitalia, 
cT. 9, Genitalia, cf , from the right. 10, Abdominal sternites 1-4, 9 . Sphaerocera tertia, 
new species: 11, Abdominal sternites and genitalia, (^. 12, Genitalia, <^, from the 
right. Sphaerocera chimborazo, new species: 13, Abdominal sternites 1-4, cf. 14, 
Abdominal sternites 1^, ? . Sphaerocera bimaculala Williston: 15, Abdominal sternites 
1-5, 9. Sphaerocera levicastilli, new species: 16, Abdominal sternites 1-4, cf. 17, 
Presternum, d^. 

Ecuador: Chimborazo, Naranjapata, Chilicay, June 16, 1955, cf 
holotype and paratype, R. Levi-Castillo, USNM type no. 66593. 

733-276— G5 2 



232 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Spliaerocera (Parasphaerocera) shannoni, new species 

Male. Resembles S. ecuadoria, new species, very closely but differs 
as follows: Legs pale yellow brown, neither tibiae nor apices of femora, 
appreciably darker. Abdominal pale spots somewhat smaller. Proster- 
nmn consisting of a very small transversely triangular plate with a 
narrow but not quite linear projection forward for a moderate distance. 
Legs, especially femora, considerably thickened, hindtibiae with no 
anteroventral spur. Male genitalia large, ninth tergite not emarginate 
to receive the cerci, which are obliquely truncate at the apex and 
separated by a deep U-shaped emargination; forceps yellow, bristly, 
and curved upward as in S. ecuadoria, rest of genitalia concealed. 
Sternites 1+2 (fig. 18) consisting of two small dots and a small 
transversely oval plate; sternite 3 broadly subtriangular but with 
posterior margin rounded, anterior margin and sides somewhat concave 
(especially in the type) ; sternite 4 trapeziform, anterior and the con- 
siderably shorter posterior margin straight, sides concave, the sternite 
twice as wide as the third ; sternite 5 somewhat asymmetrical, strongly 
transverse. 

Venezuela: San Esteban, Jan. 1-6, 1940, holotype cf , P. Anduze. 
Panama Canal Zone: Barro Colorado Island, July 9, 1923, paratype 
cf^, R. C. Shannon, USNM type no. 66594. 

Sphaerocera {jParasphaerocera) amphoraf new species 

Female. Resembles S. ecuadoria, new species, very closely but 
differs as follows: Legs whitish yellow, apical third of femora and all 
tibiae somewhat darker. Prostcrnum with a very narrow transverse 
plate attached to the mesosternum, anterior extension so narrow and 
feeble as to be scarcely visible. Legs rather long, femora slightly 
thickened, anteroventral spur on hindtibiae rather strong. Pale spot 
on abdomen slightly larger, particularly in the longitudinal direction. 
Sternites 1+2 (fig. 19) represented by a very small, distinctly trans- 
verse sclerotized spot; sternite 3 absent, but a minute black spot is 
just visible; sternite 4 elongate, somewhat narrowed at each end, 
widest three-quarters of length from anterior margin, nearly four 
times as long as broad; sternite 5 rather widely crescentic; cerci each 
with two long, almost straight bristles; 4 spiracles visible in the very 
wide, white abdominal membrane. 

Ecuador: Chimborazo, Linje, July 1955, 9 holotype; Bugna, 9 
paratype, R. Levi-Castillo, USNM type no. 66596. 

.^ Sphaerocera (Paraspliaerocerd) xiphosternum, new species 

Female. Resembles S. ecuadoria, new species, very closely but 
differs as follows: On the average, slightly smaller. Legs whitish 
yellow, apical third of femora and all tibiae somewhat darker. Pro- 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 233 

sternum with a very transverse obtusely triangular plate attached to 
the mesosternum, anterior extension short and narrow but not really 
linear except in one specimen from Higuito. Legs slightly thickened, 
hindtibiae with a very small anteroventral spur. Pale spots on ab- 
domen about the same. Sternites 1+2 (fig. 20) represented by a 
very small circular black sclerotized dot; sternite 3 normally repre- 
sented by pale membrane only, but by a small elongate dot in speci- 
men from Albrook Field; sternite 4 rather elongate, considerably 
narrowed at each end, widest at three-quarters of length from anterior 
margin, about three times as long as broad or, in the Panama speci- 
mens, somewhat less; shape of posterior end varying in Costa Rica 
specimens from rounded truncate to rounded pointed; in Panama 
specimens the sternite is considerably wider posteriorly and more 
narrowed anteriorly; sternite 5 rounded crescentic, not very strongly 
transverse; cerci each with two rather short, straight bristles. 

Costa Rica: San Mateo, Higuito, holotype and paratype 9, Pablo 
Schild. Paratypes: Costa Rica, 9, Schild (determined to be S. 
bimaculata Wilhston by Malloch). Panama Canal Zone: Albrook 
Field, June 19, 1952, 9, F. S. Blanton. Panama: Darien Province, 
Patino Point, July 12, 1952, 9, F. S. Blanton, USNM type no. 66595. 

This set of five females is somewhat variable but perhaps aU may 
belong to one species. The female from Albrook Field has a large 
mite attached to the underside of the mouth parts. 

Sphaerocera (Parasphaerocera) musiphila, new species 

Male. Resembles S. ecuadoria, new species, very closely but differs 
as follows: Legs yellowish white, apical third of femora and all tibiae 
slightly darker, femora strongly thickened, anteroventral spur of 
hindtibia very small. Prosternum with a transverse triangular plate 
attached to the mesosternum, anterior extension of moderate length, 
almost Unear. Sternites 1+2 (fig. 21) represented by a strongly 
transverse central dot and lateral dots that also are transverse; 
sternite 3 trapeziform, almost as broad as long, anterior margin 
straight, sides converging posteriorly, slightly concave, posterior 
margin also slightly concave; sternite 4 transverse, more than twice as 
wide as long, trapeziform with posterior margin shallowly emarginate, 
sides straight, diverging posteriorly; sternite 5 almost rectangular, 
much larger than 4, about fom' times as wide as long; genitalia with 
ninth tergite somewhat hollowed out on each side, emarginate to 
receive the cerci, which are separated by a wide V-shaped 
emargination. 

Panama: bananas, Sept. 8, 1932, holotype cf, CAS. 



234 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Sphaerocera (JParasphaerocera) transversalis, new species 

Male. Resembles S. ecuadoria, new species, very closely but differs 
as follows: Size slightly smaller. Facial plate (fig. 23) considerably 
shorter in longitudinal direction, more transverse. Legs yellowish 
white, apical third of femora and all tibiae slightly darker, femora 
scarcely thickened, anteroventral spur of hindtibia hardly developed. 
Prosternum at siu-face entirely membranous. Abdominal pale spots 
distinctly larger, tergite 5 with a narrow white membranous area 
both anteriorly and posteriorly. Sternites 1+2 (fig. 22) represented 
by a rounded rectangular plate that is just transverse, no lateral 
dots; sternite 3 rectangular with posterior corners slightly rounded, 
about one and one-half times as long as broad; sternite 4 rather 
small, rectangular, not quite twice as wide as long; sternite 5 partly 
concealed beneath 4, not much wider; genitalia with ninth tergite 
bearing a small central knob, somewhat emarginate to receive the 
cerci, which are separated by a U-shaped emargination. 

Ecuador: Guaj^as, Cuatro Hermanitos Experhnental Farm, Feb. 28, 
1955, holotype cf , R. Levi-Castillo, USNM type no. 66598. 

Sphaerocera (Parasphaerocera) lepidof new species 

Female. Resembles S. ecuadoria, new species, but difTers as follows: 
Facial plate more transverse. Legs yellowish white, apical third of 
femora and four hindtibiae somewhat darker, foretibiae blackish, 
anteroventral spm* of hindtibia very small. Prosternmn (fig. 25) 
shaped somewhat like a spearhead, sides angularly widened anteriorly 
where there is a narrow forward projection. Abdominal pale spots 
very large, tergite 5 divided centrally by a pale hne. Sternites 
1+2 (fig. 24) represented by an almost square central plate, not very 
small; sternite 3 considerably smaller, longer than broad; sternite 4 
narrowed anteriorly, straight posteriorly, about twice as long as 
wide; sternite 5 almost rectangular, sides hardly rounded, cerci 
concealed. 

Panama Canal Zone: Fort Kobb, Camaron, July 17, 1952, holo- 
type 9, F. S. Blanton, USNM type no. 66597. 

Sphaerocera {Parasphaerocera) transversa, new species 

Female. Close to S. lepida, new species, but evidently distinct. Legs 
whitish yellow, apices of femora slightly darker; hindtibia with no 
anteroventral spur. Abdominal pale spots relatively small, anterior 
one with sides straight, posterior margin slightly convex, posterior 
one oval with lateral ends pointed. Sternites 1+2 represented 
by a rectangular transverse plate nearly twice as wide as hindfemm-; 
sternite 4 somewhat more than twice as long as wide, posterior margin 
straight, sides slightly convergent anteriorly, anterior margin rounded; 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 235 

sternite 5 trapeziform, sides straight, as long as anterior width. 
Panama: bananas, Sept. 11, 1932, holotype 9, CAS. 

Sphaerocera (JParasphaerocera.) dissecta, new species 

Female. Allied to S. bimaculata Williston but evidently distinct. Legs 
whitish yellow; hindtibia with no anteroventral spur. Facial plate 
almost an equilateral triangle. Abdominal pale spots very large, 
square; tergite 5 consisting of two small oval plates separated by 
white membrane. Sternites 1+2 represented by a diamond-shaped 
spot not quite as wide as hindfemur, angles du'ected anteroposteriorly, 
other two laterally; sternite 3 small, width slightly greater than that 
of hindtibia, anterior margin rounded, posterior margin straight; 
sternite 4 more than twice as long as wide posteriorly, anterior margin 
rounded, posterior margin straight, sides somewhat convergent 
forward; sternite 5 rounded, crescentic, transverse. 

Panama: bananas, Sept. 22, 1932, holotype 9, CAS. 

Other Species of Subgenus Parasphaerocera 

I have placed near S. bimaculata WilHston those species of the 
subgenus that have both pale abdominal spots and entirely pale legs. 
There are a number of other species that have either the femora more 
or less blackened or the abdominal spots very reduced or absent. 

Sphaerocera (Parasphaerocera) varipes Malloch 

Sphaerocera varipes Malloch, 1925, Proc. Ent. Soc. Washington, vol. 27, p. 121. 

Malloch described this species from 5 females from Costa Rica, San 
Mateo, Higuito, and I have examined the holotype. As far as I 
can make out, there is only one species in which the femora are half 
black. This is redescribed below: 

Males and females. Black; antennae and legs yellow brown, 
coxae and proximal half of femora black, trochanters pale. Abdom- 
inal pale spots somewhat larger than in S. ecuadoria, new species, 
their margins rather rounded. Length 2.5-3.0 mm. 

Facial plate flattened, transversely trapeziform. Prosternum con- 
sisting of a small transverse triangular plate attached to the meso- 
sternum, anterior extension of moderate length, not quite linear. 
Femora distinctly thickened in the male, hindtibia with no antero- 
ventral spur. Three anterior abdominal spiracles forming distinct 
black spots in the white sternal membrane. Male with sternites 1 +2 
(fig. 26) represented by a small oval central dot; sternite 3 rather 
smaU, about twice as broad, somewhat rounded at each end and slightly 
narrowed posteriorly; sternite 4 slightly broader than long, subrec- 
tangular, posterior margin straight, sides and anterior margin slightly 



236 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iis 

rounded; sternite 5 short, very transv erse; genitalia rather small, ninth 
tergite emarginate to receive the cerci, v>^hich apparently are separated 
by a rather narrow V-shaped membranous area. Female with sternites 
1+2 (fig. 27) represented by a small oval central plate; sternite 3 not 
sclerotized; sternite 4 almost parallel sided, slightly narrowed pos- 
teriorly, where it is somewhat rounded, about three times as long as 
broad; sternite 5 somewhat widely crcscentic, cerci hidden. 

Specimens examined: Costa Rica: San Mateo, Higuito, 9 type, P. 
vSchUd; San Jose, July, 9, H. Schmidt, USNM. Panama: bananas, 
Aug. 25 and Sept. 22, 1932, 2 d", CAS. 

In the collection of the USNM are eight more females that seem to 
belong to the same species; they come from Costa Rica, Honduras, 
San Francisco ex Guatemala, and one definitely from the United 
States — Georgia: Savannah, Oct. 12, 1953, privy trap, J. W. 
Kilpatrick. 

Sphaerocera (Parasphaerocera) pallipes Malloch 

Sphaerocera pallipes Malloch, 1914, Ent. News, vol. 25, p. 31. 

This species was described originally from Panama: Buena Ventura, 
May 19, 1911, 9, USNM. I have examined the type. The species 
and its allies resemble those of the subgenus Parasphaerocera but lack 
or, in one form described below, almost lack the pale abdominal spots, 
the tergites being uniformly dark and sclerotized. Malloch (1925, 
p. 121) states he has seen many additional specimens from Panama, 
but, as I have now seen three forms (at least two species) of this group, 
I feel that the material should be re-examined. I did not find a long 
series in the USNM. 

Redescription of type: Female. Legs yellow brown, hind tibia with- 
out anteroventral spm\ Size somewhat smaller than S. ecuadoria, new 
species. Sternites 1 +2 represented by a plate that is rather narrowly 
pear shaped; sternite 3 narrow, almost linear; sternite 4 long trapezi- 
form, narrowing posteriorly, about twice as long as anteriorly broad; 
corners not rounded. Abdomen entirely black dorsally. 

Sphaerocera (Parasphaerocera) guttula, new species 

Female. Black; antennae pale brown; legs pale yeliov/ brown. 
Abdomen with very small transverse pale spots; one-fifth as wide as 
tergites, length about equal to width of hindtibia. Length about 2.5 mm. 

Facial plate almost flat, trapeziform. Prosternum with a small 
transverse triangular posterior area that is produced anteriorly into 
a short gradually narrowing extension. Hindtibia with no antero- 
ventral spur, femora slightly thickened. Tergite 5 very transverse 
posteriorly rounded. Sternites 1+2 (fig. 28) represented by a nearly 
square plate, width not quite twice that of hindfemur; sternite 3 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 237 



O 




/Su::^ 



21 




C3 22 



20 



o 






25 



26 



24 







i 







28 



27 



Figures 18-28. — Sphaerocera shannoni, new species: 18, Abdominal sternites 1-5, cf. 
Sphaerocera amphora, new species: 19, Abdominal sternites 1-S, 9 . Sphaerocera 
xiphosiernum, new species: 20, Abdominal sternites 1-S, 9 , sternite 4 shown for each of 
three specimens: a, Higuito; b, Albrook Field; c, Patino Point. Sphaerocera musiphila, 
new species: 21, Abdominal sternites \-A, cf. Sphaerocera transversalis, new species: 
22, Abdominal sternites 1-4, cf . 23, Facial plate, cf. Sphaerocera lepida, new species: 
24, Abdominal sternites 1-5, 9 . 25, Presternum, 9 . Sphaerocera varipes Malloch: 26, 
Abdominal sternites 1-4, d". 27, Abdominal sternites 1-4, 9 , sternite 3 completely 
desclerotized. Sphaerocera guttula, new species: 28, Abdominal sternites 1-4, 9- 

almost oval but posterior margin somewhat truncate, length two- 
thirds, width one-half that of sternites H-2; sternite 4 nearly square, 
much like sternites 1-1-2; sternite 5 trapeziform, sides straight, con- 
verging posteriorly, anterior and posterior margins nearly straight; 
cerci each with about three straight moderately long hairs. 

Panama Canal Zone: Erwin Island, July 18, 1923, holotype 9, 
R. C. Shannon (received in exchange from Mr. Curtis Sabrosky 
as iS*. pallipes Malloch, now in the British Museum). 



238 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Sphaerocera (JParasphaerocera) annulicornis Malloch 

Sphaeroccra annulicornis Malloch, 1913, Proc. U.S. Nat. Mus., vol. 44, p. 363. 

I have one female of this species (Missouri: Shrewsbury, Aug. 17, 
1949, W. Downes, received from Mr. Curtis W. Sabrosky). It seems 
to agree with Malloch's description. 

Female. Black; antennae brown; legs black, apices of femora, 
tibiae, and tarsi somewhat paler. Abdomen with two small trans- 
verse, laterally pointed pale spots, about half as wide as the tergites, 
and posterior one as long as width of hindfemur, anterior one not so 
long. Length nearly 3.0 mm. 

Facial plate nearly fiat, trapeziform. Hindtibia with no spur, 
femora slightly thickened. Prosternum with a very transverse 
posterior area, anterior extension short, almost linear. Tergite 5 
separated from 4 by a wide white membrane, transverse, forming a 
somewhat curved, hoodlike cover to the fifth sternite; sternites 1+2 
very transverse, subcrescentic, about half as wide as abdomen, 
bearing two longish bristles; sternites 3 and 4 not sclerotized; sternite 
5 transversely subcrescentic, fitting closely against the tergite. 

Sphaerocera iParasphaerocera) flavicoxa Malloch 

Sphaerocera flavicoxa Malloch, 1925, Proc. Ent. Soc. Washington, vol. 27, p. 121. 

I have one male paratype of this species (Costa Rica: San Mateo, 
Higuito, Pablo Schild, received through Mr. Sabrosky). It is allied 
to S. pallipes Malloch, but the facial plate is much more transverse. 

Female. Black; antennae brown, legs pale yellow brown. Abdo- 
men with two very small transverse pale spots, the first hardly 
traceable, the second one-fifth the width of the abdomen and hardly 
as long as basal width of hindtibia. Length somewhat more than 
2.0 mm. 

Facial plate flat, forming a very transverse small triangle with its 
sides somewhat concave. Hindtibia with a very small black antero- 
ventral spur, femora slightly thickened. Prosternum with a trans- 
verse posterior area and a short linear anterior extension. Fifth 
tergite transverse, posterior margin nearly straight. Sternites 1+2 
represented by dots at extreme sides and a transversely oval central 
plate about as wide as hindfemur; sternite 3 subtrapeziform with 
anterior margin straight, sides slightly concave, posterior margin 
somewhat rounded, half as wide again as 1+2, transverse, about one 
and one-half times as broad in the middle as long; sternite 4 sub- 
rectangular, nearly twice as broad as long; sternite 5 slightly visible, 
very transverse; genitalia rather large, tergite 9 not emarginate 
to receive the cerci, which are separated by a narrow U-shaped 
emargination. 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 239 

Key to Species of Subgenus Parasphaerocera Spuler ^ 

1. Abdomen dorsally with no pale spots or with very small ones, much less than 

half as long or half as wide as a tergal plate 2 

Abdomen dorsally with two large pale spots, at least two-thirds as wide or 
as long as a tergal plate 5 

2. Legs mainly black. Neither sternite 3 nor 4 sclorotized. Abdomen with 

two small pale spots (U.S.A.) . . Sphaeracera annulicornis Malloch, 1913, 9 
Legs entirely pale. Sternites 3 and 4 represented by at least small plates . 3 

3. Facial plate in form of a small, very transverse triangle. Abdomen with 

two small pale spots. Sternites (cT) 1 + 2 with a small oval central plate, 
sternite 3 transverse, rounded posteriorly, sternite 4 subrectangular (Costa 

Rica and, apparently, Brazil) S. flavicoxa Malloch, 1925 

Facial plate trapeziform, not so transverse (only 9 seen) 4 

4. Abdomen with two small pale spots. Sternites 1 + 2 large, subrectan- 

gular, sternite 3 smaller, oval, sternite 4 square, sternite 5 trapeziform 

(Panama) S, guttula, new species 

Abdomen entirely black dorsally, Sternites 1 + 2 small, pyriform, sternite 
3 narrow, sublinear, sternite 4 trapeziform, narrowed posteriorly, longer 
than broad (Panama) S. pallipes Malloch, 1913 

5. Femora with at least the basal half black 6 

Femora pale yellow, at most the distal part slightly browner or, in one 

species, the foretibiae dark 8 

6. Femora with distal half sharply pale. Sternite 3 small, pyriform, twice as 

long as broad in cf, not sclerotized in ?; sternite 4 subrectangular, in 
cf rather wider than long, in ? three times as long as broad (Central 

America and U.S.A.) S. varipes Malloch, 1913 

Femora black, apices narrowly pale (not seen, separation after Malloch) . 7 

7. Venter of 9 with two sclerotized plates. Cheeks closely striate in middle 

(Florida) S. striata Malloch, 1925 

Venter of 9 with three sclerotized plates. Cheeks not striate in middle 
(Maryland) S. nigrifeniur Malloch, 1925 

8. Males 9 

Females 15 

9. Sternite 4 not transverse, sternite 5 concealed. Ninth tergite emarginate, 

cerci separated by deep V-shaped emargination. Prosternum with 

small triangular plate with a narrow anterior projection (Ecuador) . . 10 

Sternite 4 distinctly transverse 11 

10, Facial plate raised hito a rounded convex boss. Sternite 3 parallel sided. 

S. chirtiborazo, new species 
Facial plate fiat. Sternite 3 clearly narrowed posteriorly. 

S. levicastilli, new species 

11, Sternite 3 transverse, widely trapeziform with posterior margin slightly 

concave; sternite 4 about six times as wide as long (Ecuador), 

S. trapezina, new species 
Sternite 3 not transverse 12 

12, Sternites 3 and 4 rectangular (Ecuador) . . . S. transversalis, new species 
Sternite 3 clearly not rectangular 13 

3 S. galapagensis Curran, 1934, runs down in this key to S. varipes Malloch and 
cannot be distinguished from it by any character mentioned in the description. 



240 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

13. Sternite 3 trapeziform, about as long as posteriorly broad, sides and pos- 

terior margin slightly concave (Panama) . . . S. musiphila, new species 
Sternite 3 broadly subtriangular, rounded posteriorly, one-half to one- 
third as wide as sternite 4 14 

Sternite 3 very small, not more than one-quarter as wide as sternite 4, 
elongate with concave sides, dumbbell shaped; sternite 4 five or six 
times as wide as long (Panama, Ecuador) . . . . S. tertia, new species 

14. Sternite 4 about four times as wide as long; sides of sternite 3 strongly 

concave, sternite 2 small (Ecuador) S. ecuadoria, new species 

Sternite 4 about two and one-half to three times as wide as long. Sides of 
sternite 3 weakly concave, sternite 2 much narrower (Panama and Ven- 
ezuela) S. shannoni, new species 

15. Facial plate with a rounded convex boss. Prosternum with a posterior 

triangular plate and a disconnected linear piece. Posterior pale abdominal 
spot with a linear extension to the margin. Sternites 1 + 2 relatively 
large, sternite 3 not sclerotized, sternite 4 small and elongate, sternite 

5 angular, trapeziform (Ecuador) S. eliimborazo, new species 

Facial plate flat 16 

16. Sternite 3 completely unsclerotized. Sternite 5 more or less rounded 

crescentic 17 

Sternite 3 with a distinct, even if small, plate 18 

17. Prosternum anteriorly narrow but not strictly linear. Sternite 4 smaller, 

clearly narrowed posteriorly; sternites 1 + 2 a circular dot; sternite 3 
absent, or, rarely, represented by a minute sclerotized dot (Panama, 

Costa Rica, Mexico) S. xiphosternum, new species 

Prosternum with a very transverse posterior sclerotization, anteriorly not 
visible above surface. Sternite 4 longer and less distinctly narrowed 
posteriorly; sternites 1 + 2 transverse (Ecuador). S. amphora, new species 

18. Sternite 4 long, fully four times as long as broad; sternite 5 rounded cres- 

centic (Ecuador) S. ecuadoria, new species 

Sternite 4 not very long, at most three times as long as broad 19 

19. Sternite 2 large, rectangular; sternite 5 angular, sides straight 20 

Sternite 2 dotlike or, if slightly larger, diamond shaped; sternite 5 rounded 

crescentic 21 

20. Facial plate unusually transverse. Prosternum large subtriangular. 

Foretibia black. Abdominal pale spots very large. Sternite 4 rectangular 
with anterior end almost pointed (Panama) . . . S. lepida, new species 
Facial plate normal as in figure 7. Foretibia pale. Abdominal pale spots 
smaller than usual. Sternite 4 subrectangular, rounded and somewhat 
narrowed anteriorly (Panama) S. transversa, new species 

21. Abdominal pale spots square, very large; tergite 5 divided centrally by a 

pale line. Sternite 2 diamond shaped, angles pointing anteriorly and 
posteriorly, nearly as wide as hindfemur; sternite 3 with posterior margin 
straight, anterior end rounded, not quite as wide as hindtibia; sternite 
4 with posterior margin straight, sides considerably convergent anteriorly 

where it is rounded (Panama) S. dissecta, new species 

Abdominal pale spots rounded; tergite 5 not divided by a pale line. Stergite 
2 dotlike or rectangular. Prosternum with a short very transverse piece 
posteriorly and a narrow anterior extension 22 



SPHAEROCERA IN CENTRAL AND SOUTH AMERICA — RICHARDS 241 

22. Sternite 3 dotlike, sternite 4 pyriform. Anterior part of prosternum wider. 
Abdominal pale spot as S. ecuadoria, new species (Ecuador). 

S. trapezina, new species 

Sternite 3 small but rectangular, sternite 4 rectangular, nearly three times 

as long as broad. Abdominal pale spots larger (St. Vincent and Grenada 

B.W.I.) S. bimacuiata, Williston 

The Association of Acarina with Subgenus Parasphaerocera 

Four of the males and six of the females of S. (P.) ecuadoria had 
a large mite hanging beneath the mentum. All the mites were 
attached with their heads by the foramen of the fly and with their 
dorsum directed downward. One of the mites was determined by 
Mr. E. Lindgren as Macrocheles sp. aff . insignitus Berlese. A specimen 
of S. (P.) xiphosternum from Panama, Albrook Field, had a similar 
mite on its mentum. 

Literature Cited 

Malloch, J. R. 

1913. Description of new species of American flies of the family Bor- 

boridae. Proc. U.S. Nat. Mus., vol. 44, no. 195S, pp. 361-372. 

1914. A new Borborid (Diptera) from Panama. Ent. News, vol. 25, p. 31. 
1925. A synopsis of New World flies of the genus Sphaerocera (Diptera; 

Borboridae). Proc. Ent. Soc. Washington, vol. 27, pp. 117-123. 
Richards, O. W. 

1961. Diptera (Sphaeroceridae) from South Chile. Proc. Roy. Ent. Soc. 
London, ser. B, vol. 30, pp. 37-68, 13 figs. 
Spuler, a. 

1924. North American species of the genus Sphaerocera and Aptilotus (Dip- 
tera-Borboridae). Pan-Pacific Ent., vol. 1, pp. 66-77, 4 figs. 
Williston, S. W. 

1896. On the Diptera of St. Vincent (West Indies) . Trans. Ent. Soc. London, 
pp. 253-446. 
Young, Benjamin P. 

1921. Attachment of the abdomen to the thorax in Diptera. Cornell 
Univ. Agric. Exp. Sta. Mem., no. 44, pp. 255-306, illustr. 



U.S. COVERNMENT PRINTING OFFICEll96B 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, B.C. 

Volume 116 1965 Number 3505 



HERPETOLOGY 

OF THE ZUNI MOUNTAINS REGION, 

NORTHWESTERN NEW MEXICO^ 



By Frederick R. Gehlbach^ 



Introduction 



In decided contrast to the amount of herpetological information 
available for other southwestern states, relatively little is known 
about amphibians and reptiles in New Mexico. Several taxonomic 
papers have dealt with members of the state herpetofauna, but none 
have treated local distribution and differentiation in detail. It 
is the intent of this study, and of others in progress, to describe the 
amphibians and reptiles of New Mexico by natural regions, with the 
aim of understanding local variation in morphology, distribution, 
and life history. 

Field observations and specimens obtained during portions of the 
summers of 1951, 1954, and 1955 constituted a prelimiuar}'^ survey of 
amphibian and reptilian species in northwestern New Mexico. 
Thereafter, emphasis was placed on the Zuni region as a biogeographic 
unit. Local studies of ecology and assumed gene flow were made 

1 Modified from a master's thesis submitted to Cornell University, 1959. 

2 Department of Biology, Baylor University, Waco, Texas. 

243 



244 PROCEEDINGS OF THE NATIONAL MUSEUM vol. hg 

intermittently during June-September in 1956 and 1957, and during 
June-August in 1958 and 1959. Additional data on vegetation were 
recorded in August 1960; a few herpetological specimens were added 
at this time and in July-August 1961. Altogether, 132 days and 
nights were spent in the field in the area covered by figure 1 . 

A long history of random herpetological collecting in the Zuni 
region has preceded these investigations. S. W. Woodhouse, a 
member of the 1851 Sitgreaves Expedition, first obtained repre- 
sentatives of the local herpetofauna. Two new species, Tapaya 
oniatissima {—Phrynosoma douglassi hernandesi) and Pityophis affinis 
{=Pituophis melanoleucus affinis) were described from among his 
specimens. Personnel associated with the United States Army at 
Fort Wingate, the U.S. Biological Survey, and various U.S. railway 
and army explorations west of the one-hundredth meridian, especially 
V. Bailey, E. A. Goldman, H. W. Henshaw, C. G. Newberry, and 
R. W. Shufeldt, contributed specimens prior to 1910. This histori- 
cally important material is housed in the U.S. National Museum. 
E. T. Hooper and W. L. Chenoweth collected amphibians and reptiles 
in 1939 and 1949-1951 respectively; their specimens and my own 
have been deposited in the Museum of Zoology, University of Michigan, 
and the Cornell University herpetological collection. 

Aspects of this study were discussed with several persons, especially 
Bruce B. Collette, Norman Hartweg, Charles F. Walker, and Thomas 
M. Uzzell, Jr. William J. Hamilton, Jr., at Cornell University and 
Norman Hartweg and Charles F. Walker at the University of Michigan 
greatly facilitated my research. William L. Chenoweth, Hillis L. 
Howie, and Rossiter D. Olmstead were contributors to success in the 
field. Edward C. Raney provided the opportunity for a trip to the 
U.S. National Museum, where Doris M. Cochran aided my quest 
for specimens and historical data. Stanwyn G. Shetler identified 
certain plants. My wife, Nancy Y. Gehlbach, helped with vegeta- 
tional analyses and typed the manuscript. A Sigma Xi grant-in-aid 
for research made part of the 1958 field work possible. 

The following loaned specimens or provided specific information 
about them: Curtis Allen, James D. Anderson, Ralph W. Axtell, 
W. Frank Blair, J, Roger Bider, Charles M. Bogert, James E. Bohlke, 
Bryce C. Brown, William G. Degenhardt, Floyd L. Downs, William 
E. Duellman, James S. Findley, Joe Gorman, Michael Hays, Arthur 
H. Harris, Norman Hartweg, Robert F. Inger, Clyde J. Jones, Laurence 
M. Klauber, John M. Legler, Paul Licht, Edmond V. Malnate, 
T. Paul MasHn, Samuel B. McDowell, Robert R. Miller, Jon A. 
Peterson, Paul W. Parmalee, Jay M. Sheppard, Hobart M. Smith, 
Wilmer W. Tanner, Robert W. Risebrough, Robert C. Stebbins, 
David A. West, Ernest E. WilUams, Kenneth L. WiUiams, and 
Richard G. Zweifel. 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 245 

I am very grateful to each of the above persons and institutions. 
I am equally indebted to the Prairie Trekers, whose efforts in my 
behalf produced many specimens and much inspiration; I dedicate 
this paper to them. 

Physical and Climatic Environments 

The study area, referred to as the Zuni region or Zunis, includes 
most of McKinley and Valencia Counties, New Mexico (fig. 1). It 
is situated in portions of both Datil and Navajo Sections of the 
Colorado Plateau at the southeastern extremity of this physiographic 
province (Hunt, 1956, fig. 1). Elevations range from 5794 to 11,389 
feet. The diverse topography and semiarid climate, with their 
attendant effects on vegetation, greatly influence herpetofaunal 
distribution. 

Topography 

Most spacious of the various topographical features are the Zuni 
Mountains, a domal uplift approximately 25 miles wide and 70 miles 
long, lying northwest by southeast near the southeastern edge of 
the Colorado Plateau. Strata surrounding the dome are gently 
sloping Pennsylvanian through Cretaceous formations, mostly sand- 
stones (Fenneman, 1931, pp. 217-319). Rock outcrops are exposed 
in canyons, mesas, and the great retreating cliffs characteristic of 
Colorado Plateau topography. These major erosional features 
probably were formed no earlier than the Miocene and, along with 
the entire Plateau, have been uplifted in Pliocene and Pleistocene 
times (Hunt, 1956). The summit of Mount Sedgwick at 9156 feet 
is the highest point above the basal plain, which averages 7000 feet. 

Running along Oso Ridge, the Continental Divide separates the 
Zuni Mountains almost equally into eastern and western watersheds. 
The former is partly drained by Azul and Bluewater Creeks, which 
flow into the Rio San Jose and ultimately into the Rio Grande. The 
latter is drained by the Rio Puerco (this name is also given to a stream 
of the Rio Grande basin), Rio Nutria, and Zuni River, which empty 
into the Little Colorado River. All streams, except the headwaters 
of the Zuni, are intermittent, and steady flow occm"s only during the 
summer period of heaviest precipitation. Bluewater Lake and 
Ramah Reservoir are relatively permanent man-made features. 

Between the Zuni Mountains and other structural upwarps are 
broad, open valleys such as the Gallup-Zuni Basin. The strata of 
this basin slope gently westward from the Zuni Mountains, decreasing 
from 8000 feet to 6500 feet toward the mouth of the Zuni River. 
Such features give the region its continuous yet vertically diverse 
aspect. To the north, gently undulating topography of the Navajo 



PROCEEDINGS OF THE NATIONAL MUSEUM 




HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 247 

Section continues into the San Juan Basin. On tlie south the Zunis 
and North Plains of the Datil Section are replaced by the San Augus- 
tine Plains and rugged montane highlands associated with the Basin 
and Range physiographic province. 

East of the Zuni Mountains, Mount Taylor, an 11,389 foot Miocene 
volcano, dominates the area of eroded, basalt-capped mesas known 
collectively as the San Mateo Mountains (not to be confused with an 
uplift of the same name in Socorro County). Mesa Chivato is the 
northeastern extension of this highland mass, which is approximately 
15 by 35 miles in extent. Its slopes are drained by the Rio San Jose 
and Chico Arroyo and drop to 5794 feet at Laguna on the eastern 
end of the study area. This is about the southeastern boundary of 
the Colorado Plateau as marked by the westernmost faults of the 
Rio Grande (Hunt, 1956, p. 6). 

Extensive lava beds or malpais, derived in part from Mount Taylor, 
are most conspicuous south of Grants. They stretch 30 miles on a 
southwesterl3^-northeasterly axis and are surrounded by a relatively 
flat plain several feet lower than the overrun lava. Cinder cones 
border the main malpais, which ranges from 8308 feet at the top of 
Flagpole Crater to 6200 feet near McCarte3^s. Much soil has ac- 
cumulated where the lava is relatively smooth, as in the vicinity of 
El Morro National Monument and Cebolleta Mesa. This basalt is 
probably of late Miocene origin, whereas rough, broken malpais along 
the Rio San Jose near Grants is of Pleistocene derivation (Hunt, 1956, 
pp. 46, 53). The McCarteys lava may be less than 1200 3^ears old 
(Nichols, 1946, p. 1049). North of Bluewater another small malpais 
lies isolated on the Cretaceous plain and, near it, numerous volcanic 
necks stud the landscape. 

Climate 

The Zunis are characterized by a semiarid-mesothermal climatic 
regime with a P-E Index of 16-32 (Thorn thwaite, 1931). It may be 
seen that local moisture conditions are allied closely with elevation 
and exposure (fig. 2). Prevailing air currents are southwesterly. 
Below 8000 feet average annual precipitation values are 13 inches on 
the southwest and 10 on the northeast; above 8000 feet, 20 inches is 
the average annual value (U.S. Dept. Comm., 1945-1957). The 
period of greatest rainfall occurs from July through September. 
Short, heavy thundershowers are then common and often result in 
flash floods. Heavy winter snows usually are limited to the elevations 
above 8000 feet. 

Pearson (1931, table 14) has assembled annual mean precipitation 
data for various plant zones in Arizona and New Mexico. They 
range from 11 inches in the grasslands and 16 in pinyon-juniper 



248 



PROCEEDINGS OF THE NATIONAL MUSEUM 



woodlands to 21 iii ponderosa pine and 22 in Douglas fir zones. The 
evaporative effect of winds, however, Unaits mesophytic vegetation 
on exposed slopes so that correlations between precipitation and 
elevation are often spurious for indexing vegetational distribution. 

80 r 



< 
(r 

UJ 
Q. 

:e 

UJ 



70 



60 

hi 
o 

LlJ 

^ 50 



40 



30 



20 



> 8 



SAN FIDEL 6100 ft. 

EL MORRO 7218ft. 




nr. McGAFFY 8200 ft. 



1.0 L4 L8 2.2 2.6 

PRECIPITATION inches 



3.0 



Figure 2. — Cllmatograph of three weather stations in the Zuni region. (Plotted points 
are mean monthly temperatures and precipitation; months are indicated by numbers. 
Data compiled from U.S. Department of Commerce climatological summaries for New 
Mexico, 1945-1957.) 

Air temperatures generally fall with increasing elevation, the annual 
fluctuation being equally great at all stations (fig. 2). The number 
of days between 32° F. extremes is 123-194 (x 162) below 7000 feet, 
72-148 (120) above 7000 feet (U.S. Dept. Comm., 1945-1957). 
Daily temperature variation is also considerable (table 1). Pearson 
(1931, tables 3, 4) calculated June-September maximum and mini- 
mum temperatures in New Mexico and Arizona as follows: 90, 55 
(°F) in the grassland zone; 83, 52, pinyon-juniper woodland; 79, 46, 
ponderosa pine zone; and 74, 40, in the Douglas fir zone. Hooper 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



249 



(1941, table 1) and Lindsey (1951, table 1) have presented other 
temperature data for the Zuni region. 

Table 1. — Temperature and humidity in the pinyon-juniper-ponderosa pine 
ecotone, 7200 feet, Cottonwood Gulch, McKinley County, New Mexico 



Measurement 


1959 


Value 




Avg. max. temp. 


July 1-31 


93° F. 






August 1-20 


87 




Avg. min. temp. 


July 1-31 


51 






August 1-20 


47 




Max. temp. 


July 3 


98 




Min. temp. 


July 3 


38 




Avg. rel. humidity 








7:00 A.M. 


July 1-August 10 


56% 




1:00 P.M. 


July 1-August 10 


29% 





Ecological Distribution 

Included m the Navahonian biotic province of Dice (1943), the 
Zuni region contains Plains, Roughlands, and Montane Life Belts. 
While these biogeographic units vary with ecologic succession and 
exposure, they may be employed to describe elevational distribution 
of amphibians and reptiles. Within each Hfe belt, the characteriza- 
tion of vegetational associations and substrate conditions will further 
delimit herpetofaunal range. No attempt at complete vegetational 
analysis is made here; only the relative positions of certain aspect 
dominants, important in providing cover, are described. The Hfe 
belts and associations are both vertically and horizontally discon- 
tinuous, becoming increasingly mesophytic mth ascending elevation. 

Plains Life Belt 

This lowermost unit extends in patchwork fashion from 5794 feet at 
the eastern end of the study area to approximately 7200 feet on the 
lava beds near El Alorro. Shrub cover is relatively sparse except 
in the vicinity of arroyos and around rock outcrops and mesas. The 
soil may consist of loose sand or clay with Uttle rock or restricted 
areas of gravel and rubble. At the edges of the Zuni and San Mateo 
Mountains this Hfe belt is broken by mesas included in the Rough- 
lands Life Belt and by malpais. Part of Bailey's (1913, pp. 25-41) 
Upper Sonoran Zone, the mixed grasslands of Castetter (1956), the 
basin sagebrush and short-grass plains of Clements (1920, pp. 139- 
144, 156, 160); and Pearson's (1931) grassland zone are included in 
the Plams Life Belt. 



250 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Eight (27 percent) of the 30 species of amphibians and reptiles 
recorded from the Zuni region are, so far as known, restricted to this 
belt. They are Scaphiopus intermontanus , Scaphiopus bombifrons, 
Bujo pundatus, Uta stansburiana, Holbrookia maculata, Thamnophis 
dorsalis, Masticophis jiagellum, and Crotalus atrox. Certain other 
species, especially Sceloporus graciosus, Eumeces obsoletus, Pituophis 
melanoleucus, and Crotalus viridis, appear to reach their maximum 
abundance here. 

For the most part, the Plains Life Belt is characterized by the 
short-grass or mixed -grass association in which blue grama (Bouteloua 
gracilis) is dominant or codominant with such other grasses as hairy 
grama (B. hirsuta) and galleta {Hilar ia jamesi) (pi. lA). Holbrookia 
maculata usually is found where shrub cover is sparse and composed 
of rabbitbush {Chrysothamnus nauseosus), snakeweed (Gutierrezia 
sarothrae), or narrow-leaved yucca (Yucca glauca). Reptile life is 
more varied in somewhat heavier cover, as in the saltbush-sage 
association dominated by shadscale saltbush (Atriplex canescens), 
spiny saltbush (A. conjertijolia) , or big sagebrush {Artemisia triden- 
tata). Sceloporus graciosus is Umited primarily to this vegetational 
type. Crotalus atrox seems similarly limited to the cholla-juniper 
association dominated by cholla cactus (Opuntia imbricata) and 
one-seed juniper {Juniperus monosperma) . 

Between elevations of 6500 and 7200 feet there may be an ecotone or 
a broad continuum between Plains and Roughlands Life Belts marked 
by an increased number of one-seed junipers and various taller shrubs 
such as mountain mahogany (Cercocarpus montanus). Pinyons 
(Pinus edulus) may be present. The widespread continuum could, 
in part, be caused by excessive gi'azing (Castetter, 1956, p. 272). 
Eumeces obsoletus occure here as do most other Plains species that 
range into the Roughlands. Rock-strewn arroyos also permit species 
like Crotaphytus collaris and Masticophis taeniatus to bridge the 
ecological gap between the two life belts. 

Roughlands Life Belt 

The Roughlands Life Belt comprises rocky uplands from around 
6800 feet to at least 8000 feet. Its outside elevational limits are 
influenced by the presence of rocky soil combined with the effects of 
less moisture at lower elevations and lower temperatures in Montane 
environments (see Castetter, 1956, p. 271). Vegetative cover is much 
thicker than on the plains, and outcropping sedimentary strata produce 
boulder piles and talus slopes, thus providing additional retreats for 
secretive species. This life belt includes the upper part of Bailey's 
(1913, pp. 25-46) Upper Sonoran Zone and lower limits of his Tran- 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 251 

sition Zone in addition to the woodland biome of Castetter (1956), 
pinyon-cedar woodland and petran chaparral of Clements (1920, pp. 
183-187, 197-199), and pmyon-jimiper and lower ponderosa pine 
zones of Pearson (1931). 

Two species, Diadophis punctatus and Salvadora gmhamiae, pres- 
ently are known only from this life belt but probably are more 
widely distributed. Ambystoma tigrinum, Scaphiopus hammondi, 
Bufo woodhousei, Rana pipiens, Hyla arenicolor, Phrynosoma douglassi, 
Sceloporus undulatus, Urosaurus ornatus, Eumeces multivirgatus, 
Cnemidophorus velox, and Thamnophis elegans reach then* gi-eatest 
abundance here. Other, rarer species, such as Hypsiglena torquata 
and Crotalus molossus, may be centered in the Roughlands Life Belt. 

Dominated by pinyons, one-seed junipers, alligator junipers (Juni- 
perus deppeana), and Rocky Mountams junipers (J. scopulorum) , the 
pinyon-juniper association is the most widespread vegetational type 
(pi. IB). Shrub cover is relatively sparse but consists of aspect 
dominants similar to those of the oak-mahogany association de- 
scribed below. Amphibians and reptiles such as Scaphiopus ham- 
mondi, Phrynosoma douglassi, and Cnemidophorus velox are more 
abundant where the pinyon-juniper resembles a rocky savanna. 
Openings in the savanna often are occupied by saltbush and occa- 
sionally extensive stands of big sagebrush. If large boulders are 
present, as on hillsides and mesa edges, Crotaphytus collaris and 
Urosaurus ornatus are typically present. 

The oak-mahogany association is composed largely of Gambel oak 
(Quercus gambeli) and mountain mahogany. Below approximately 
7000 feet, wavyleaf oak (Q. undulata) may replace Gambel oak. 
Shrubs like gooseberry {Ribes sp.), skunkbush (Rhus trilobata), and 
datil (Yucca baccata) are present in varying numbers. This associa- 
tion occupies canyon sides and may replace the pinyon-juniper or 
ponderosa pine where timber is cut. Marked by heavier, more mesic 
cover, it shelters Bufo woodhousei, Eumeces midtivirgatus, and some 
adults of Ambystoma tigrinum. It is distinct from the riparian associ- 
ation of canyon bottoms that is dominated by the narrowleaf cotton- 
wood (Populus angustijolia) and various willows (Salix spp.) (pi. IB). 
Hyla arenicolor, Rana pipiens, and Thamnophis elegans are more or 
less limited to the vicinity of streams and ephemeral ponds resulting 
from stream subsidence in the riparian association. 

Transition from pinyon-juniper woodland into the Alontane Life 
Belt often is gi-adual, especially on the north side of the Zuni Moun- 
tains and on Mount Taylor. Ponderosa pine (Pinus ponderosa) 
occurs in a few, relatively pure stands as low as 7200 feet. In the 
ponderosa pine association, mountain muhly (Muhlenbergia montana) 
may be the predominant ground cover. Pituophis melanoleucus and 



252 PROCEEDINGS OF THE NATIONAL MUSEUM vol. hg 

Crotalus viridis were collected here as were representatives of all 
species found to be more abundant in pinyoji-juniper, oak-niohagany, 
and riparian associations. 

Montane Life Belt 

The heavy cover characteristic of this uppermost life belt prevails 
above approximately 7800 feet; lower limits may be found in sheltered 
canyons and on north-facing slopes, where exposure is a restricting 
factor. The ponderosa pine association marks the lower boundary 
and may include Douglas fir (Pseudotsuga menziesi) in cooler situa- 
tions. Upper limits are best developed on the higher peaks, where 
the spruce-fir association includes Engelmann spruce {Picea engel- 
manni) and Douglas fir along with alpine fir (Abies lasiocarpa) and 
other conifers (pi. IC). With the exception of parks and the limited 
alpine tundra association on Mount Taylor, vegetation is usually too 
thick to permit much heating of the substrate. The soil is composed 
of humus covered by conifer needles. Bailey's (1913, pp. 41-51) 
Transition, Canadian, and Hudsonian Life Zones are included in this 
belt as are the coniferous forest and petran alpine biomes of Castetter 
(1956), petran montane and subalpine forests of Clements (1920, pp. 
207-210, 224-226), and Pearson's (1931) ponderosa pine, Douglas 
fir, and Englemann spruce zones. 

No reptiles or amphibians are restricted to this belt and none appear 
to reach their maximum abundance here. While 14 species were 
collected in montane associations, all of the reptiles except Thamnophis 
elegans came from logged-off areas or rocky openings in ponderosa 
pine and spruce-fu' associations. Rana pipiens, Bi/fo woodhousei, 
and Hyla arenicolor, among others, follow the riparian association 
that also penetrates the Montane Life Belt. Ambystoma tigrinum, 
Pseudacris triseriata, Eumeces multivirgatus, Phrynosoma douglassi, 
and T. elegans were fouiid in parks or stands of quaking aspen 
{Populus tremuloides) (pi. IC). Only P. douglassi was taken in the 
alpine tundra association, which has no arboreal vegetation. 

Recent Environmental Changes 

There is little doubt that the Zuni region has suffered considerable 
modification within the past hundred years. The impact of man is 
superimposed on a record of continuous climatic change. Drought 
reduced the water supply and, coupled with grazing pressure, has 
eliminated much of the original grassland (Castetter, 1956, pp. 269, 
272). Statements on the former abundance of grass were made by 
Bigelow (1856, p. 6), Whipple (1856, p. 62), and many others. Eoth- 
rock (1875, p. 120) described open parklike stands of ponderosa pine 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 253 

with good forage at about 8000 feet near Ft. Wingate and noted that 
fir covered surrounding hillsides. Woodlands are now much thicker 
in some places and thinner or nonexistant in others because of grazing 
and logging, respectively (Castetter, 1956, pp. 272, 277), Correlated 
changes in horpetofaunal distribution may be siu-mized. 

Surface water was more abundant prior to the twentieth century, 
Whipple (1856, pp. 14, 63) noted that the Zuni Indians cultivated 
without hrigation and described Agua Fria as a permanent spring. 
Kennerly (1856, pp. 9-10) observed the Zuni River as a small, clear 
stream in which fishes were secm'cd at several localities. He also 
noted maay frogs along the Rio Pescado ia November. Coues 
(1875, pp. 614-615) found Thamnophis elegans in the Zuni River, 
"wherever this stream spread into sluggish lagoons." While seasonal 
changes influenced differences of opinion as to the abundance of 
water (e.g., in September, Sitgreaves, 1854, p. 5, described the Zuni 
River as a mere rivulet), it is noteworthy that Agua Fria and the 
lower Rio Pescado and Zuni are presently intermittent. Zuni Indians 
now irrigate; streams in the vicinity of their pueblo are extensively 
controlled. 

As suggested by extensive arroyo cutting, which began in the late 
1880's (Bryan, 1925, 1928), drought may account for the disappear- 
ance of surface water not purposely diverted by man. When this 
study was initiated, the Zuni region, indeed the entire Southwest, was 
m the midst of severe drought. Breeding amphibians were not 
collected until July 1954 and remained uncommon until 1957. Diado- 
phis punctatus was not discovered at a previously worked locality 
(Cottonwood Gulch) until August 1959. The drought ended locally 
in 1957 when the Colorado Plateau Section of New Mexico received 
146 percent of average annual moisture (U.S. Dept. Comm., 1945- 
1957). Profound changes in the neighboiing Texas herpetofauna 
were linked to this drought (Blair, 1957; Milstead, 1960). 

Iq 1960 the Bureau of Sport Fisheries exterminated all native 
aquatic organisms in the upper Zuni River (R. R, Miller, in litt.). 
Not a single aquatic animal depending on dissolved oxygen was left; 
an apparently undescribed fish, genus Pantosteus, was among the forms 
eliminated. Such wanton destruction probably will continue as the 
human population continues to increase. Grants experienced a 
354 percent rise in population between 1950 and 1960 (Grants Cham- 
ber of Commerce, in litt.). Fortunately, however, small portions of 
Zuni grassland and piny on- juniper association are preserved in EI 
Morro National Monument and hopefully will be preserved in the 
proposed Manuelito National Monument. 



254 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iis 

Lava 

In contrast to numerous investigations of coloration in lavicolous 
mammals, relatively few workers have shown simUar interest in 
lava-dwelling amphibians and reptUes. Lewis (1949; 1951), Norris 
(1958, p. 270), and Lawrence and Wilhoft (1958) have demonstrated, 
however, the presence of abnormally dark reptiles on southwestern 
lava flows. Apparently selection favoring procrypsis is operating in 
these environments. Tertiary lava beds in the Zuni region were 
described adequately by Hooper (1941) and Lindsey (1951). Hooper 
found that mammals restricted to the malpais had not developed dark 
races there. 

With the exception of Ufa stansburiana, Sceloporus graciosus, 
Masticophis Jlagellum, and Crotalus atrox, the species characteristic 
of the Plains Life Belt occasionally were taken on soil-covered lava. 
This substrate, widespread near El Morro and CeboUeta Mesa, pre- 
sents an appearance similar to the short-grass association except that 
it is generally higher in elevation (see Lindsey, 1951, figs. 10, 11). 
The Grants lava flow, on the other hand, is rough and relatively 
unweathered, with less soil accumulation (see Hooper, 1941, pi. 2). 
Crotaphytus collaris, Sceloporus undulatus, Urosaurus ornatus, Eumeces 
obsoletus, Thamnophis elegans, Pituophis melanoleucus, and Crotalus 
viridis were collected or observed on the rougher malpais. 

Only C. collaris, U. ornatus, and Scaphiopus hammondi show 
evidence of unusually dark coloration. The latter two species are 
predominantly dark gray with little trace of the usual patterned 
dorsum. Because local populations of these species tend to resemble 
substrate color to a remarkable extent, it appears unlikely that the 
dark, lavicolous individuals represent unique variation. Lewis (1951) 
found dark U. ornatus in southern New Mexico. Three adults of C. 
collaris are patterned but are quite dark in general coloration. This 
is the usual situation on New Mexico lava flows (Fitch, 1956a, p. 223). 

Numbers of lavicolous reptiles and amphibians were not large. 
While the rubble of broken lava offers uncountable hiding places, the 
presence of ponderosa pine and Douglas fir, postclimax to adjacent 
nonlava vegetation (Lindsey, 1951, p. 220), may limit the ranges of 
Plains species on such a substrate. Sections of sparsely vegetated 
malpais dominated by apache plume (Fallugia paradoxa) held the most 
reptiles, but there appeared to be easy access to and from the lava 
margins, with the possibility of strong immigration pressure from 
contiguous grassland supporting larger populations of several species. 
Gene flow may thus reduce the effectiveness of local selective pres- 
sures. Marginal habitats, relative accessibility, and geologic re- 
cency of the Grants malpais undoubtedly account for the fact that no 
indigenous dark races have developed there. 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 255 

Vertical Distribution 

In comparison with the Guadalupe Mountains, a region of about 
equal size and vertical range in southeastern New Mexico, it is evident 
that Zuni environmental diversity exceeds the diversity of the Zuni 
herpetofauna. Only 7 (33 percent) of 21 Zuni genera contain more 
than one species whereas 17 (41 percent) of 42 genera in the Guada- 
lupes have two or more species (Gehlbach, in ms.). Sixty-five am- 
phibians and reptiles currently are recorded in the Guadalupe region; 
only 30 definitely are known in the Zunis. This relative paucity may 
be influenced directly by the high elevations and climatic extremes. 
With increasing elevation there is a decline in the number of species 
(fig. 3) . Such forms as Scaphiopus homhijrons, Eumeces obsoletus, and 
Crotalus molossus, near the boundary of their range in the Zunis, are 
restricted locally in contrast to their greater vertical distributions 
elsewhere. 

Vertical transects from Thoreau, McEanley County, south to 
Cottonwood Gulch, thence to Lookout Mountain and El Morro, 
Valencia County (7000-9100 feet), and from Grants to Mount Taylor, 
Valencia County (6400-11,389 feet), were repeated in several sum- 
mers (fig. 1). These, the locality records, and miscellaneous observa- 
tions demonstrate that no two morphologically related species have 
precisely the same vertical range or relative abundance (fig. 3) . This 
displacement pattern is especially apparent within genera but also 
occurs between genera if species possess similar habits (e.g., between 
Holbrookia maculata and Phrytiosoma douglassi, or between Masticophis 
taeniatus and Pituophis melanoleucus) . Axtell (1959, table 1) and 
Lowe and Zweifel (1952, fig. 4) have diagramed similar patterns in 
west Texas and central New Mexico respectively. The probability 
that two species will not be equally efficient in the same environment 
has been discussed by many biologists (e.g., Lack, 1949; Hutchinson, 
1957). 

Ecological differences were observed most readily among the Zuni 
lizards (table 2). Since similar differences have been correlated with 
morphological variation (Collette, 1961; Lundelius, 1957) and thermo- 
regulation (Bogert, 1949; Fitch, 1956b), the presence or absence of 
such correlations may help to elucidate local ecological segi'egation. 
Sceloporus undulaius, S. graciosus, Urosaurus ornatus, Phrynosoma 
douglassi, Eumeces multivirgatus, and Cnemidophorus velox are of 
similar size; hence, presumably they are able to capture similar-sized 
prey. Attention is focused on these species with particular reference 
to the number of lamellae as an index to climbing ability (Collette, 
1961). 

Of the iguanids, U. ornatus shows the greatest climbing propensities 
and has the most lamellae in proportion to size (cf. table 2; fig. 4). 



256 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Ambystoma tigrinum 



Scaphiopus hammondi 




Bufo woodhousei 
Hyla arenicolor 
Rano pipiens 
Crotaphytus collaris 
Holbrookia maculata 
Phrynosoma douglassi 
Sceloporus graciosus 
Sceloporus undulatus 
Urosaurus ornatus 



■11,300 




Eumeces multivirgatus 
Eumeces obsoletus 
Cnemidophorus velox 
Masticophis taeniatus 
Pituophis melanoieucus 
Thamnophis elegans 
Crofalus atrox 



Crotalus viridis 

I I I I I I I I I 

5800 62 66 70 74 78 82 86 9000 

ELEVATION-feet 

Figure 3. — Schematic vertical distribution and relative abundance of some amphibians 
and reptiles in the Zuni region. (The horizontal line indicates marginal occurrence; the 
widest part of each polygon is the approximate elev^ation of greatest abundance irrespec- 
tive of habitat considerations noted in the text.) 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 257 

Its elevational range is about equal to that of S. undulatus, but the 
two species are separated ecologically in all life belts. As suggested 
by its fewer lamellae, S. undulatus is primarily terrestrial, while U. 
ornaius is distinctly scansorial and saxicolous. At the same locality 
S. undulatus is found on the ground or on fallen or leaning tree trunks 
(rarely on vertical trunks), and U. ornatus inhabits rock outcrops, 
boulder piles, and cliffs. U. ornatus maintains body temperatures 
that are two to three degrees higher than S. undulatus or S. graciosus 
(P. Licht, in litt.). It frequents situations with greater exposure 
(table 2). 

Table 2. — Niche relationships of adult lizards in the Zuni region {species are 
arranged from most scansorial to most terrestrial, top to bottom of table) 



Shade 


Partial shade 


Sun 






Urosaurus ornatus 




Sceloporus -undulatus 


Crotaphytus collaris 




Sceloporus graciosus 


Cnemidophorus velox 




Eumeces obsolctus 


Holbrookia maculata 


Eumeces multivirgatns 


Phrynosoma doxtglassi 





S. graciosus also is predominantly terrestrial and does not occur 
in ecological sympatry with U. ornatus. It has more lamellae than 
S. undulatus and fewer than U. ornatus; its vertical range and center 
of abundance are quite different (fig. 3). S. graciosus was found 
only in the vicinity of big sagebrush or saltbush on loose soil. In 
contrast, S. undulatus rarely was seen on such a substrate and was 
taken commonly on hard-packed clay or broken sandstone in the 
pinyon-juniper savanna. While these edaphic factors may separate 
partially the two species of Sceloporus, marked terrestrialism in S. 
graciosus belies the obvious adaptive significance of having more 
lamellae. This character actually may have little to do with local 
ecological segregation, or it could permit more effective locomotion 
on loose soils. 

P. douglassi has the lowest lamellae count among the iguanids con- 
sidered here and is characteristically the most terrestrial (table 2 ; fig. 
4). Although the two species of Sceloporus sometimes climb in rock- 
piles, bushes, and trees, P. douglassi was never observed off the ground. 
It has the greatest elevational range of any species in the Zunis and is 
more sedentary in habit and more abundant above 7200 feet (fig. 3). 
The vertical distribution and lamellae count of E. multivirgatus are 
most like those of P. douglassi ; however, this skink typically inhabits 
dense vegetative cover (table 2). Bogert (1949) and Fitch (1956b) 
have shown that low body temperatures are more typical of Eumeces 
than either Sceloporus or Cnemidophorus. 



258 



PROCEEDINGS OF THE NATIONAL MUSEUM 



The very high lamellae count of C. velox apparently is unrelated to 
scansorial ability, for this teiid rarely climbs. Instead, might the 
lamellae count be correlated with a longer toe, as Hecht (1952, p. 116) 
has suggested in other lizards? The number of lamellae cannot be 
used as an index of relative climbing ability here; its inapplicability 

90 



E 80|- 

X 



:z 

UJ 

> 

I 

O 



C/) 



60 



50 



40 





=,,,,0, 



10 



15 



20 



25 



30 



35 



LAMELLAE 



Figure 4. — Relationship of the number of fourth-toe lamellae to snout-vent length in 
adults of some Zuni lizards. (Outside limits of the plotted points are joined to form 
polygons representing each species. Means are indicated by position of the symbols.) 



I Phrynosoma douglassi 
A Eumeces multwirgatus 
A Sceloporus undulatus 



X Sceloporus graciosus 
9 Urosaurus ornatus 
O Cnemidophorus velox 



with respect to the observed inbergeneric differences is apparent 
(fig. 4). Ecological separation is indicated, however; C. velox in- 
habits areas of greater insolation than the other nonclimbing species 
(table 2). Its congeners maintain much higher body temperatures 
than the other genera studied (Bogert, 1949; Fitch, 1956b). 



PROC. U.S. NAT. MUS. VOL. 116 



GEHLBACH — PLATE 1 




C2>l%?-^ 



\ Short-grass association of the Plains Life Belt near Laguna, Wilencia County, New 
'Mexico, ca. 5700 feet, June 1957; B, Roughlands Life Belt at Cottonwood Gulch, 
McKinlev Countv, New Mexico, ca. 7200 feet, June 1957 (the hillside supports pinyon- 
juniper association with riparian association on the right); C. Montane Life Belt at Ojo 
Redondo, Valencia County, New Mexico, ca. 8600 feet, July 1954 (a natural park in the 
foreground is surrounded by spruce-fir association perforated by aspen stands). 



PROC. U.S. NAT. MUS. VOL. 116 



GEHLBACH — PLATE 2 







^-3 ^ 



O ^ 



U 



£ S o 

I— ' . <~o 







H-. o 
O 3 g 

■5b "5 S 



3 '^ ^^ 

U '— I .« 

■5 N " 

g S a, 

o ^ " 






W 



^ 



PROC U.S. NAT. MUS, VOL. 116 



GEHLBACH — PLATE 3 











m 



i 



B 



A, Ontogenetic series of C«^m:c^oMo'-M^ "^^/o-v (UMMZ 120288-89, 120309) from the Zun, 
region: snout-vent lengths from left to right are 80, 70, 61, and 36 mm.; note the 
absence of spots, also the complete mlddorsal light stripe on the left followed by a 
specimen lacking this stripe and two with incomplete stripes. B, Ontogenetic series of 
Cnemidophorus Ixsau^uis (UMMZ 79212, 15052, 102729-30) from Grant County, New 
Mexico: snout-vent lengths from left to right are 87, 81, 65. and 39 mm.; note the grad- 
ual appearance of spots. 



PROC. US. NAT. MUS. VOL. 116 



GEHLBACH — PLATE 4 




Z, M 



o o 



U 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 259 

Accounts of Species 

Adult specimens of Zuni amphibians and reptiles that have been 
criticaUy examined number about 700. Many more larval amphibians 
and collections from outside areas also have been studied. Meristic, 
morphometric, and color-pattern variations are analyzed according 
to taxonomic criteria set forth by the latest revisers. These criteria 
are recognizably different, depending on the student of the group 
(Pimentel, 1958). Their value, in certain cases, may be challenged. 
In addition, novel characters are described and evaluated with 
particular regard for environmental and ontogenetic variation. 

For most species, a brief, locally pertinent, nomenclatorial history 
is followed by taxonomic analysis, data on life history, and a summary 
of ecological distribution. A more penetrating taxonomic account 
is included for Ambystoma tigrinum, Cnemidophorus velox, and Diado- 
phis pundaius in view of certain problems that exist. Less space is 
devoted to some species because other workers are utihzmg Zuni 
material or because few specimens or observations actually are 

available. , • i tv/t + 

Locality records refer to the Zuni herpetofauna exclusively. Most 
of these records are based on specimens examined, but a few, indicated 
by asterisks, represent museum data suppHed by colleagues. Com- 
parative material examined includes only those extralimital specimens 
used in figures and tables or specifically referred to in the separate 
accounts and not given full provenance data in these discussions. 
In some cases newer maps permit more accurate mileages and ele- 
vations than previously published by Chenoweth (1950), Gehlbach 
(1956), and Hooper (1941); however, the Zuni region remains to be 
thoroughly surveyed. . . 

Museum specimens are denoted by the following abbreviations: 

AMNH American Museum of Natural History, New York, N.Y. 

ANSP Academy of Natural Sciences, Philadelphia, Pa. 

BCB Bryce C. Brown (private collection), Waco, Tex. 

BYU Brigham Young University, Provo, Utah 

CCNP Carlsbad Caverns National Park, Carlsbad, N.M. 

CNHM Chicago Natural History Museum, Chicago, 111. 

CU Cornell University, Ithaca, N.Y. 

CWGF Cottonwood Gulch Foundation, Thoreau, N.M. 

ISM lUinois State Museum, Springfield, 111. 

KUMNH Kansas University, Museum of Natural History, Lawrence, Kans. 

LMK Laurence M. Klauber (private collection), San Diego, Calif. 

MCZ Museum of Comparative Zoology, Harvard Univ., Cambridge, 

Mass. 

MVZ Museum of Vertebrate Zoology, Univ. of California, Berkeley, 

Calif. 
TNHC University of Texas Natural History Collection, Austin, Tex. 
UCM University of Colorado Museum, Boulder, Colo. 
738-287—65 2 



260 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

UIMNH University of Illinois, Museum of Natural History, Urbana, 111. 
UMMZ University of Michigan, Museum of Zoologj', Ann Arbor, Mich. 
UNMCV University of New Mexico, Collection of Vertebrates, Albuquer- 
que, N.M. 
USNM United States National Museum, Washington, D.C. 

Class Amphibia: Order Caudata 

Ambystoma tigrinum nebulosum Hallowell 

The subspecific status of tiger salamanders on the Colorado Plateau 
and in adjacent areas has been interpreted variously. Allocation of 
Zuni specimens was left in doubt by Dunn (1940, p. 162), Bishop 
(1943) and Stebbins (1951; 1954) included the Zunis within the 
range of A. t. nebulosum, possibly because some of Dunn's specimens 
resembled material from Utah and Colorado then known as A. t. 
nebulosum through confusion of type locality. Lowe (1955d, p. 244) 
correctly reassigned the name nebulosum to A. tigrinum of the 
Mogollon Plateau and vicinity. But as evidence now indicates, 
Lowe (1955d, p. 247) inadvertently confused the status of Colorado 
Plateau populations by describing A. t. utahense from this region. 

Adult specimens, above 80 mm. snout-vent length, of A. tigrinum 
from the Zunis have relatively longer tails in larger individuals, 
especially the males (fig. 5) . With two exceptions, their color pattern 
consists of yellow spots or reticulations on a brownish to black ground 
(pi. 2C). Five adults have reticulate patterns and five subadults, 
60-73 mm. snout-vent, resemble them (pi. 2D). In 15 other adults 
the number of dorsal spots between extremes of limb insertions ranges 
from 11 to 50 (x 30.8±2.8); tail spots number 8 to 37 (13.0±2.9). 
These resemble A. t. stebbinsi on the basis of dorsal spots but are closer 
to A. t. nebulosum as regards tail spots (cf. Lowe, 1955d). When 
compared to the topotypic A. t. nebulosum described by Lowe (1955d, 
table 1), the Zuni salamanders average more dorsal body spots and 
fewer tail blotches. Their venters are mottled as in A. t. nebulosum 
or variously suffused with dark pigment as in A. t. stebbinsi or A. t. 
utahense (pi. 2B; cf. Lowe, 1955d, figs. 4A, C, 5A). They are as 
distinct from topotypic A. t. nebulosum as A. t. stebbinsi is and com- 
bine the diagnostic features of both races. I suggest that A. t. steb- 
binsi belongs in the synonymy of A. t nebulosum. 

In contrast to the specimens described above, one adult male 
(KUMNH 45762, 95 mm. snout-vent) is much darker in general 
aspect, with dorsal spots reduced to minute points of dull yellow, 
although the tail spots are plainly indicated. Another large, dark 
male is olive brown with irregular black body markings and light- 
olive tail spots (pi. 2A). These specimens resemble A. t. utahense or 
A. t. utahense x nebulosum, as does a similar specimen from San 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



261 



Francisco Mountain, Coconino County, Arizona, the type locality of 
A. t. nebulosum (see Hallowell, 1854, p. 144). 

Evidence from other Colorado Plateau locaHties coupled with the 
Zuni data suggests that A. t. utahense is an ontogenetic variant of 
A t nebulosum. Lowe's statement (1955d, p. 238) that adult color 



90 



80 



^70 
E 
X 
5 60 



< 50 



40 



30 



30 



A-4V* 



A e 



40 






o •© 



o° o 



o 
o o o 



50 60 70 80 90 

SNOUT-VENT LENGTH mm. 



100 



Figure 5.-Relationship of tail length to snout-vent length in Ambystoma tigrinum nebu- 
losum from the Zuni region. 



# Males, metamorphosed 

y Females, metamorphosed |Yellow-spotted 

O Larvae J 

■ Males, metamorphosed] Non-yellow-spotted 

A Metamorphosed jg^p^^.^^^^^jly ^ai^t^ined (collected July 16, 1957, 7800 feet) 

A Larvae J 



pattern is established upon transformation, unfortunately, is mis- 
leading. Dunn (1940, p. 154) and Bishop (1941, p. 163) have sum- 
marized ontogenetic changes in color pattern with which I concur. 
In fact, Dunn (1940, p. 158) diagnosed A. t. nebulosum {=A. t 
utahense, Lowe, 1955d) as containing yeUow-spotted young and 



262 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

non-yellow-spotted adults. It is significant that recently trans- 
formed Zuni specimens, like certain larvae, are olive brown above 
with small scattered dark spots and some lemon-yellow mottling on 
the tail, thus approximating A. t. nehulosum figured by Bishop (1943, 
fig. 45: 1,2). Only later do they develop the yellow spotting charac- 
teristic of subadults. 

Two individuals from Molas Lake, San Juan County, Colorado, 
kept alive for over two years, have undergone continuous color- 
pattern change. At metamorphosis (ca. 65 mm. snout-vent) they 
resembled A. t. utahense in the same manner as the Zuni specimens. 
Within a week the ground color darkened and yellow spots appeared. 
Other specimens (UMMZ 121925) preserved at this time were also 
like A. t. nehulosum. The yellow spots then enlarged and became dull, 
changing to olive; the original ground color was reduced to small, 
irregular black marks. One (UMMZ 122526) died in May 1961; it 
measures 101 mm. snout-vent. The other presently resembles UMMZ 
120301 (see pi. 2A) from the Zunis and is 96 mm. snout-vent. A 
single live individual from Telluride, San Miguel County, Colorado, 
resembled A. t. nehulosum at 70 mm. but currently is closest to A. t. 
utahense at 105 mm. 

Dark-brown, black-spotted adults taken in ponds near Telluride 
(UMMZ 120304) and at Molas Lake (UMMZ 121926-29) are 81- 
98 mm. snout-vent, in contrast to smaller, 63-69 mm., yellow-spotted 
individuals taken with them. A similar size difference occurs in 
"utahens e-\ike" adults, 92-107 mm., and "nehulosum-like" subadults, 
68-73 mm., from the Chuska Mountains, San Juan County, New 
Mexico (UNMCV 1111-21). I have also examined small yellow- 
spotted and reticulated specimens from Coconino County, Ai'izona 
(UMMZ 120302), and San Juan County, Utah (UMMZ 121931). 
This rough correlation of size and coloration favors the hypothesis 
of ontogenetic change. Sexual pattern dimorphism is absent; how- 
ever, relative size at metamorphosis may influence color pattern 
(see below). Since A. t. utahense as defined by Lowe (1955d, pp. 
246-247) is based apparently on ontogenetic variation in A. t. nehu- 
losum, it is suggested that this name also be placed in the synon- 
ymy of ^. t. nehulosum. 

Color-pattern comparisons of larvae, 35-65 mm. snout-vent, from 
the Colorado Plateau and adjacent areas reveal no basic geographic 
differences. The coloration described by Stebbins (1951, p. 46), 
however, may vary locally as regards density of spotting and shade of 
ground color; for example, the specimens from Catron County, New 
Mexico (UNMCV 1 163-68), are heavUy spotted or mottled with brown 
and black. The tail fin has black edges, and the lateral light line or 
row of light spots is nearly indistinct. In contrast, specimens from 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



263 



Bernalillo County (UNMCV 1138-61) are very light with little hint of 
spotting. It is apparent that larvae from shallow, weU-illuminated 
pools are lighter in color than those from deeper, darker environments. 
Very large larvae are usually greenish, olive, or brown, with widely 
spaced black spots. They often lack light and dark mottling and a 
lateral light line. Coues (1875, p. 633) described this "Siredon" or 
axolotl coloration in specimens from Jacob's Well, Apache County, 
Arizona, and from San Francisco Mountain. 

The nrnnber of gill rakers provides structural evidence that larvae 
from the Colorado Plateau, southern Arizona, Wyoming, and other 
localities in New Mexico are very similar (table 3). Northern Arizona 
specimens have somewhat fewer gill rakers, but this appears to be local 
variation since larvae from Telluride, Colorado, resemble them yet 
are geographically closer to populations with more rakers, e.g., Molas 
Lake. Larvae from Michigan (A. t. tigrinum) and San Luis Potosi, 
Mexico (A. f. velasci), also have fewer gill rakers and approach the 
reduced number characteristic of Ambystoma rosaceum in this respect. 
Further study of larval characters should aid in clarifying the status 
of other subspecies of tiger salamanders. 



Table 3. — Frequency distribution of gill rakers on the anterior face of the third gill 
arch in larvae {35-65 mm. snout-vent) of Ambystoma tigrinum and A. rosaceum 







Gill rakers 














10 11 12 13 14 


15 16 17 


18 


19 


20 


21 


22 


23 N 


X 


A. rosaceum 




















Durango, Laguna del Progresso 


3 5 1 














9 


10.8 


Durango, 2.5 mi. W. San Luis 


2 5 2 














9 


11.0 


Sonora, El Tlgre Mountains 


2 5 2 1 














10 


12.2 


A. tigrinum 




















San Luis Potosi, San Diego 




1 6 


2 


1 








10 


17.3 


Michigan, Ann Arbor 




2 


6 


2 


1 






10 


18.3 


Arizona, Hauchuca Mountains 








2 


2 


4 


2 


10 


20.6 


New Mexico, 9 mi. E. Mogollon 








1 


2 


2 


1 


7 


20.9 


New Mexico, Zunl region 








4 


7 


2 


2 


I 16 


20.3 


New Mexico, 10 mi. S. Albuquerque 










3 


4 


2 


L 10 


21.1 


New Mexico, Sandia Park 








2 


3 


2 


1 


9 


20.5 


New Mexico, Pecos 






1 


3 


1 


4 




10 


20.2 


Arizona, Wiieatfield Creek 






1 


3 


1 


2 




7 


19.6 


Arizona, Grand Canyon 




1 


2 


3 


3 


1 




10 


19.1 


Colorado, 9 mi. SW. Telluride 






2 


2 


5 


1 




10 


19.5 


Colorado, Molas Lake 








2 


1 


6 


1 


10 


20.6 


Wyoming, Hoback Basin 








2 


2 


2 


4 


10 


20.8 



In the Zunis, larvae and adults of A. t. nebulosum usually were found 
in temporary potholes and canyon-bottom pools. Larvae were col- 
lected in cattle tanks on Mount Taylor in 1959, and S. W. Woodhouse 
(1854, p. 35) reported Siredon lichenoides (=large larvae of A. tigrinum) 
in a spring near the Zuni Pueblo. Although suitable habitats were 
investigated repeatedly throughout the present study, larvae were not 
discovered until July 1957, when the unusually heavy rainfall produced 



264 PROCEEDINGS OF THE NATIONAL MUSEUM vol. no 

great amounts of surface water. Prior to that time, one active adult 
was dug from moist ground in the riparian association at a depth of 
six feet during excavations for a swimming pool. 

One hundred and fifty-three larvae, similar in coloration to that 
described by Stebbins (1951, p. 46) from near Flagstaff, Arizona, were 
seined from a pothole approximately 10 feet in diameter on July 16, 
1957, at 7800 feet. Of these, 36 measured 35-45 (x 42.2) mm. snout- 
vent, 32-39 (35.8) mm. tail length. In the laboratory, 25 were kept 
for 28 days, 5 to a gallon jar. At the end of this period, 5 specimens 
had died, 14 had fully transformed, and 6 still possessed well-developed 
gills and larval body form. Body proportions of these specimens 
reveal greater size in the recently metamorphosed individuals (fig. 5). 
During captivity the larvae continuously gulped air at the water 
surface, something they had not been doing when collected. 

On July 27, 1957, larvae equal in size to those previously captured 
at 7800 feet were taken from an arroyo 500 feet lower at the mouth of 
the same canyon and nearly one mile distant. A week of heavy rain 
that flooded the first locality filled the previously dry arroyo and 
presumably carried larvae to the new locality. 

Five large larvae and one adult were collected in the pothole at 
7800 feet, July 3, 1958, prior to the onset of heavy summer rainfall. 
The pond was 15 feet wider at this time than previously. About 
20 other A. iigrinum of similar size could be seen surfacing and gulp- 
ing air. The majority of these had normal larval gills and body 
form, but several had reduced gills. No other adults were observed 
although several were taken in ponds later in the summer. The 
collected larvae were light olive green dorsally with scattered dark 
spots. Two transforming specimens had light lateral reticulations 
on the tail; theu" venters were unmarked. These five larvae and 
six others from a pool at 7000 feet are of similar coloration and pro- 
portions. They have relatively longer tails than transformed speci- 
mens of similar body size (fig. 5). 

The above data suggest that larval development requires at least 
two years in some populations of ^. t. nebulosum in the Zunis. First 
proposed by Burger (1950b), this life history pattern does not apply 
necessarily to all Zuni larvae, for shallow pools at 7300 feet examined 
July 3, 1958, contained small larvae similar in size to the first-year 
individuals taken in 1957. Spatial isolation of larval size groups 
is not surprising when one considers the predacious nature of these 
salamanders. The tadpoles of Hyla arenicolor and Bufo woodhousei, 
although abundant in the area, rarely were found m the same pools 
with A. tlgmium. Food habits of tiger salamander larvae during the 
anuran breeding season demonstrate why this is so (fig. 6). 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



265 



That paedogenesis occurs in larger Zuiii larvae is evidenced by 
the fact that at least one spechnen (USNM 8240, 103 mm. snout-vent) 
contains many large yolked eggs. Environmental factors believed 
to cause neoteny in A. tlgrinum have been widely reviewed. In 
the present study, transformation, both natural and experimentally 
tested, seemed to be initiated by shrinkage of the pond environment 
associated with increased temperatures and decreased oxygen supply. 



MISC. DIPTERA 
LARVAE 
ADULT AQUATIC 
COLEOPTERA 

NOTONECTID 
NYMPHS 



fHIRUDINEA 27o 

,ANISOPTERA NAIADS 2% 




Figure 6.— Food (in percent of volume) of 36 larvae of Ambysloma tigrinum nebulosum, 
35-45 mm. in snout-vent length, collected July 16, 1957, at 7800 feet. 

Shufeldt (1885) cited similar evidence for metamorphosis in Zuni 
larvae as did Durham (1956, p. 220) for A. tigrinum from northern 
Arizona. Schuierer (1958) said that, among other things, a well 
oxygenated, constant water supply was impHcated in maintaining 

neoteny. 

Most Zuni larvae forced to gulp air at the surface m laboratory 
jars metamorphosed within four weeks of capture regardless of size. 
Specimens under similar conditions remained larval in form when 
provided with an artificial oxygen supply. Some larvae, approxi- 



266 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

mately 50 mm. snout-vent, in an exposed, rapidly evaporating pool, 
transformed during a 59-day observation period in 1959. Another 
group of similar size located under a ledge in a nearby pool showed 
little reduction of larval features during this time. Thus, A. tigrinum 
in the Zunis may remain larval and become paedogenic if provided 
with a relatively permanent aquatic environment but can transform 
under arid conditions. Such ecotypic plasticity also has been noted 
in Amby stoma macrodactylum by Kezer and Farner (1955). 

Locality records: 

MCKINLEY CO.: 6 mi. S. Thoreau (CU 6234, 6303-4; UMMZ 120301, 120305- 
6); 8 mi. S. Thoreau (CU 6301-2, 6411; UMMZ 122929); 4 mi. NNE. Prewitt 
(CU6307); Ft. Wingate (USNM 14409-12, 14402, 14633, 14490, 14818, 19118); 
Nutria (USNM 8240, 8456); 2.5 mi. N. Upper Nutria (KUNMH 45762); Val- 
encia CO.: El Morro Nat. Mon. (KUMNH 15556-7); 17.5-18 mi. NE. Grants 
(UNMCV 1122-28). 

Comparative material examined: 

Ambystoma tigrinum nebulosum. — New Mexico: Bernalillo co. : Sandia 
Park (UNMCV 1129-37); 1 mi. W. and 10 mi. S. Alburquerque (UNMCV 1138- 
61); CATRON CO.: 9 ml. E. Mogollon (UNMCV 1163-68); san miguel co.: 
Pecos (UMMZ 67886, 71383); san juan co.: 5 mi. W. Toadlena (UNMCV 
1111-13); 2 mi. S. Washington Pass (UNMCV 1119-21); Arizona: apache co.: 
Wheatfield Cr., 0.5. mi. W. Ariz.-N. M. boundary (UMMZ 121932); coconino 
CO.: Greenland Lake, Grand Canyon Nat. Park (UMMZ 120302); 2. mi. N. 
North Rim entrance, Grand Canyon Nat. Park (UMMZ 120303); santa 
CRUZ CO.: Parker Canyon, Huachuca Mts. (CNHM 62616, 62638, 62640); 
Colorado: san miguel co.: 1 mi. W. Telluride (UMMZ 121943); 9 mi. SSW. 
Telluride (UMMZ 120304); san juan co.: Molas Lake (UMMZ 121923-29); 
Utah: san juan co. : Monticello (UMMZ 121931); Wyoming: sublette co.: 
Hoback Basin (UMMZ 99751). A. t. tigrinvjn. — Michigan: washtenaw co.: 
Ann Arbor (UMMZ 69648). A. t. velasci.— Mexico: san luis potosi: San 
Diego (UMMZ 77331). A. rosaceum. — Mexico: durango: Laguna del Pro- 
gresso (UMMZ 102528, 110902); 2.5 mi. W. San Luis (UMMZ 110903-4); 
sonora: Santa Maria Mine, El Tigre Mts. (UMMZ 78353).^ 

Order Anura 

Scaphiopus hanimondi Baird, S. intermontanus Cope, and 
S. bonibifrons Cope 

Members of the subgenus Spea of Scaphiopus have been treated 
differently with respect to specific status. Linsdale (1940, p. 200), 
VanDenburgh (1924, p. 194), and Wright and Wright (1949, pp. 
112-120) considered ^S. intermontanus and S. homhijrons to be races 
of S. hammondi, an arrangement first proposed by Cope (1889). 
S. intermontanus was placed as the northern subspecies of S. hammondi, 



* These hirvae were reported as A. tigrinum by Stebbins (1951, p. 40, pi. 52) 
but apparently they are A. rosaceum because of their striking hght- and dark- 
reticulated pattern (unlike any tigrinum known to me) and because of their fewer 
gill rakers (table 3). 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 267 

and S. homhijrons was considered a distinct species by Schmidt 
(1953, p. 59) and Stebbins (1951, pp. 202-218; 1954, pp. 92-94). 
Stejneger and Barbour (1943, pp. 37-39), Tanner (1939), and Zweifel 
(1956) separated the three forms as species, an arrangement which 
I favor on the basis of available Zuni material. I further concur 
with Zweifel (1956, p. 37) in suggesting that S. bombifrons and S. 
intermontanus may be conspecific. Blair (1956) thought the call 
of S. intermontanus to be more nearly like that of S. bombifrons than 

S. hammondi. 

The measurements of 41 adult spadefoots, considered to be *b. 
hammondi on the basis of flattened frontoparietals, are summarized 
(table 4). Males are somewhat larger than females as noted in 
S. holbrooki by Pearson (1955) . Such sexual dimorphism is frequently 
reversed in other anurans. Specimens from the Miocene lava plains 
near El Morro National Monument are larger and darker in color 
than those from light, sandy soils near Thoreau and Grants. When 
compared with individuals "from other southwestern states, the total 
series is considerably smaller in all respects (see Tanner, 1939, p. 17). 
All adults of S. hammondi have red-tipped dorsal tubercles, and 
most have small dark spots or blotches scattered over the dorsum. 

Four adults of S. bombifrons and two of S. intermontanus ^ are 
distinct when one considers relative elevation of the frontoparietal 
bones as the best criterion for separation (see Zweifel, 1956, pp. 
35-37). Unlike S. hammondi, these six specimens possess two 
continuous dorsolateral light stripes that diverge posterolaterally 
from the occiput. The S. bombifrons measure 48, 49, 50, and 52 mm. 
snout-vent; the S. intermontanus, 46 and 51 mm. These individuals 
are, therefore, larger than most S. hammondi from the Zunis. A 
size difference between S. hammondi and S. bomUfrons-S. intermon- 
tanus may serve as an isolating mechanism in this area of sympatry. 
Lowe (1954) has discussed this factor in Scaphiopiis of south-central 
New Mexico. 

S. intermontanus is recorded here from New Mexico for the hrst 
time. The specimens were taken at 7200 feet in an area of mixed 
grass and one-seed junipers. The present localities for S. bombifrons, 
at 6400 to 7200 feet in the Plains Life Belt, are near the extreme 
northern limit of its range in New Mexico; hence, an ideal situation 
for studying the relationships of S. bombifrons and S. intermontanus 
exists in the Zunis. Conclusive evidence that these two species are 
isolated reproductively rests with the collection of additional material 
in this critical area. 

The spawning of S. hammondi in rainpools, cattle tanks, roadside 
ditches, and canyon-bottom potholes was observed during periods of 
heavy rainfall in July and August. Breeding sites were located in 



268 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 269 

all major vegetational associations below the spruce-fir, but western 
spadefoots were most abundant in the pinyon-juniper savanna and 
Plains-Roughlands continuum. The earliest observed egg-laying 
occurred July 4, 1954, at 6400 feet in a small tank on the eastern 
soil-covered edge of the Grants malpais; the air temperatm-e was 
48° F., water 53° F., at 9:30 p.m. 

In the choruses of S. hammondi, most vocal males swam or floated 
in the middle of the pond, but several called from shallow water or 
shore. S. homhijrons and S. intermontanus were not taken in breeding 
aggregations. They may breed earlier in the season here as S. 
intermontanus does in Utah (Blair, 1956). This could explain the 
shore-caUing station of S. hammondi in view of its apparent preference 
for a swimming-floating position when in reproductive sympatry 
with other spadefoots (Gehlbach, 1956, p. 365; McAlister, 1959). 

The larvae of S. hammondi commonly occurred with tadpoles of 
Hyla arenicolor and Bufo woodhousei in the riparian association of the 
Roughlands Life Belt. They were found only with Bufo punctatus 
in choUa-juniper grassland and without anuran associates in the 
short-grass association. On July 3, 1959, cannibalistic larvae of 
S. hammondi with hyper trophied jaw muscles and beaked upper 
mandibles were coUected at 7700 feet in pinyon-juniper-ponderosa 
ecotone. They were located with typical larvae in a rainpool four 
inches deep and two by six feet in outside dimensions. The water 
temperature was 75° F., air 83° F., at 11:20 a.m. A smgle dip net 
sample held 15 cannibalistic (6-13, x 10.9 mm. body length) and 
6 typical (16-18, x 17.4 mm.) specimens, the former with hindlimb 
buds, the latter with larger hindlimbs and toes. Seven recently 
transformed Scaphiopus species, taken at 6900 feet, September 1925, 
by Neil M. Judd, measure 16-22 (x 18.6) mm. snout-vent. 

Locality records: 

S. hammondi.— ucKiNLKY co.: 6 mi. S. Thoreau (CU 6309; UMMZ 120300); 
8 mi. S. Thoreau (CU 6476; UMMZ 120308); Thoreau (USNM 70412-21;; 
16 mi. S. Gallup (CU 6231) ; Valencia cc: 15 mi. E. Grants (KUMNH 14547-66, 
14569-72); 11.5 mi. SSE. Grants (CU 5775); El Morro Nat. Mon. (KUMNH 
14538-46, 17271-8). S. bombifrons.—ucKiNhEY CO.: 16 mi. S. Gallup (CU 6310); 
VALENCIA CO.: Grants (KUMNH 14568); El Morro Nat. Mon. (KUMNH 
17222-3). S. iiitermontanus.— VALENCIA CO.: 2 mi. W. El Morro Nat. Mou. 
(CU 6308). 

Bufo punctatus Baird 

This red-spotted toad is known from only two localities in the Zuni 
region. It apparently does not breed above 6800 feet, for intensive 
search has not revealed its presence in numerous anuran choruses in 
or above the pinyon-juniper association. Thus, it is somewhat 
ecologically separated from Bufo woodhousei, which is most abundant 
above 7000 feet (fig. 3). B. punctatus may be common in the choUa- 



270 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

juniper grassland and Plains-Roughlands continuum around Grants. 
A single chorus observed at 6400 feet, July 4, 1954, after a heavy rain, 
consisted of more than 100 calling males in a small cattle tank on 
soil-covered lava adjacent to the rough Grants malpais (Gehlbach, 
1956, p. 365). Scaphiopus hammondi was present in about equal 
numbers in the breeding aggregation. 

Locality records: 

VALENCIA CO.: 11.5 mi. SSE. Grants (CU 5766); Grants (Smith, 1950, fig. 45). 

Bufo woodhousei woodhousei Girard x H. w. australis Shannon 

and Lowe 

Rocky Mountains toads in Arizona recently have been split into 
two races (Shannon and Lowe, 1955). The nominate, northern 
subspecies is now considered to occupy the Colorado Plateau in that 
state; it is separated presumably from a new southern, lowland form, 
B. w. australis, by the MogoUon Plateau. Shannon and Lowe (1955, 
pp. 188, 190) included New Mexico within the range of B. w. australis 
but did not state specific localities. B. woodhousei from the Zuni 
region does not conform readily to either subspecies. Geographically 
these toads might be expected to resemble typical B. woodhousei, 
but structurally they combine some characters of B. w. australis with 
B. w. woodhousei and are considered intergrades. 

With regard to diagnostic features, 22 adult specimens have: 
(1) frontal area of craDium elevated m 91 percent; (2) snout-vent 
length/parotoid gland width 10.6-13.9 (x 11.9±0.19); (3) snout-vent 
length/parotoid gland length 4.4-5.6 (5.1 ±0.74); (4) skin of upper 
surfaces smooth; (5) median white line on snout present in 82 percent; 
(6) black pectoral semicircles not strongly developed; (7) dorsal color 
brown with 68 percent having large warts set in black. Characters 
1, 2, 4, 5, and 6 favor allocation with B. w. woodhousei, while 3 and 7 
are closer to B. w. australis. 

Colorado Plateau populations of B. woodhousei should be more 
or less uniform in the Zunis and adjacent Arizona since no important 
barriers to east-west dispersal exist. Such continuity is shown, for 
the characters of B. w. woodhousei predominate in the specunens at 
hand. The features of B. id. australis, however, indicate that gene 
flow from the south occurs if, in fact, these features are sufficient to 
distinguish B. w. australis from the nominate form in New Mexico. 
Irrespective of the validity of the taxonomic characters involved, the 
Rio Grande valley and associated lowlands provide a suitable path- 
way for north-south gene flow in central New Mexico (see discussion 
under "Zoogeography," p. 315). 

Although Shannon and Lowe (1955) furnish ratios of parotoid gland 
size to body length, they do not limit these criteria to any certain age 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



271 



group or sex. The validity of such ratios in terms of isometric growth 
of the parotoid gland and sexual dimorphism can be tested (fig. 7). 
It may be seen that juvenile specimens have a relatively larger gland 
and that gland width is more variable than length. No sexual 
dimorphism is apparent, but females are too few to demonstrate this 
adequately. In juveniles of less than approximately 30 mm. snout- 
vent, the cranial crests often are barely discernible, and the frontal 
area of the cramium undeveloped. These featm-es and the parotoid 
gland, therefore, should be utilized to separate larger specimens only, 
preferably sexually mature adults (table 4). 



.14 



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A A 

A 



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20 30 40 50 60 70 
SNOUT- VENT LENGTH mm. 



80 



90 



100 



Figure 7.-Relationship of parotoid gland length/snout-vent length (lower symbols) and 
parotoid gland width/snout-vent length (upper symbols) to body size in Bufo woodhousei 
from the Zuni region. 

# Sexually mature males Q Sexually mature females A Juveniles 

The larvae of B. woodhousei have been collected throughout the 
summer months in slowly running streams, potholes, and cattle tanks. 
A sample of 12 taken at 7800 feet, July 3, 1959, are 7-12 (x 9.4) mm. 
body length; they have hindlimb buds. One day later, at 7100 feet, 
12 larvae with well-developed legs and complete tails were collected; 
they measure 15-17 (15.7) mm. and approximate the size of trans- 
forming B. woodhousei from northern Arizona (Wright and Wright^ 
1949, p. 209). Six recently metamorphosed individuals, 13-19 (x 
16.9) mm. snout-vent, were found on August 8, 1957, at 7000 feet. 



272 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

That the Ziini toads have a biannual breeding regime is indicated 
by the formation of a chorus July 29, 1957, in a pond where large 
larvae were already present. Choruses often were heard after the 
initial summer rains, suggesting that perhaps summer and spring 
peaks of precipitation initiate breeding. Gravid females were found 
at two such choruses, but amplexed pahs were not seen. Thornton 
(1960, p. 181) found previously marked females of B. woodhousei 
in amplexus twice in the same breeding season at Austin, Texas. 
Bragg (1941, p. 112) stated that B. woodhousei breeds more or less 
independently of rain at Las Vegas, New Mexico. 

Juveniles, previously characterized (Gehlbach, 1956, p. 365), were 
often in evidence during the day in riparian and oak-mahogany 
associations. Eight collected July 4, 1959, at 7100 feet, along with 
the transforming larvae mentioned above, measure 25-34 (x 31.2) mm. 
snout-vent. Adults were distinctly crepuscular or, on warm nights, 
nocturnal above the grasslands. They were not as common near 
arroyos in the Plains Life Belt as in more mesic situations at higher 
elevations. Their opportunistic feeding habits are indicated by speci- 
mens with stomachs containing only Phyllophaga or Eleodes species 
when these beetles were abundant. It is noteworthy that the latter 
tenebrionid genus is thought to be distasteful to many predators 
(T. Eisner, inlitt.). 

Locality records: 

MCKINLEY CO.: Thoreau (MCZ 32813*); 5.5 mi. S. Thoreau (CU 6312); 6 
mi. S. Thoreau (CU 5765, 5767, 6332); 7 mi. S. Thoreau (CU 6337); 16 mi. S. 
Gallup (CU 6313); 14 mi. N. and 7 mi. E. Gallup (UCM 6544-49); 4 mi. W. 
Page (CU 6334); Ramah Reservoir (CU 6311); 5 mi. SW. Ramah (UMMZ 
123133); Upper Nutria (UMMZ 123134); Ft. Wingate (USNM 14395, MVZ 
9342-48*); Valencia co.: 8 mi. SE. Paxton (UMMZ 86604); Grants (MVZ 
57640*); Mt. Sedgwick USNM 54363) ; 8 mi. SE. Thoreau (UMMZ 120298-99, 
120307). 

Hyla arenicolor Cope 

The taxonomic status of canyon tree frogs from the Zunis is being 
studied elsewhere; certain observations on adult size and metachrosis, 
nevertheless, seem worthy of record here. Canyon tree frogs exhibit 
considerable color-pattern variation (Stebbins, 1951, p. 315). Zuni 
specimens were spotted usually with greenish or dusky, Uchen-Uke 
markings, but when exposed to sunlight, they often became a uniform 
dirty cream or grayish white. The skin of such exposed individuals 
was noticeably more pustulose than in recently captured specimens 
found in dark crevices or cracks between rocks. The ground color 
of most H. arenicolor was yellow cream, tan, or light reddish brown, 
closely approximating the sandstone substrate on which they were 
restine-. 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 273 

Like Bufo woodkousei, this hylid may have two or more brecdijig 
seasons coinciding with separate peaks of precipitation. Heavy July 
rains initiated calling in pools where adult H. arenicolor had not been 
in evidence previously. Two adult females with large yolked eggs 
(ca. 1.6 mm. diameter) were collected July 6 and August 27, 1956; 
they measure 45 and 50 mm. snout-vent and are larger than most 
males taken with them (table 4) . 

Four transforming larvae, 17 to 20 mm. body length, were found 
July 2-6, 1956, at 7800 feet; eggs Avere laid July 16, 1956, in the same 
pond. On the latter date, in 1957, larvae in all stages including 
some with external gills were coUected at 7800 feet. The water 
temperature was 75° F. at 11:20 a.m., the ah- 83° F., at this same 
locahty on July 3, 1959, when a similar series was found. Of these 
specimens, 12 legless larvae measure 9-13 (x 10.9) mm. body length; 
12 with both pairs of legs are 14-17 (15.7); and 12 recently trans- 
formed are 22-28 (24.7) mm. snout-vent. They are similar m 
size-group composition to a series (UMMZ 121950) obtained Septem- 
ber 4, 1960, near Snowflake, Navajo County, Arizona, by Robert E. 
Miller, and they are larger than larvae from the Chuicahua Mountams, 
Cochise County, Arizona (Zweifel, 1961). 

Adults and larvae were found often in rocky canyon pools of the 
riparian association at elevations where the pinyon- juniper or ponder- 
osa pine associations dominated surrounding hillsides (pi. IB). The 
tadpoles were usually associated mth larvae of Bwfo woodhousei and 
Rana pipiens in potholes 1-10 feet in diameter and not more than 10 
inches deep. Such habitats were devoid of aquatic vegetation and 
probably resulted from recent stream subsidence. On July 18, 1956, 
five calling males were found in water-filled, eroded pits on the top of a 
sandstone outcrop at 6400 feet. This outcrop was isolated by level 
grassland, characterized by junipers and scattered pinyons at the east 
edge of the Grants malpais. 
Locality records: 

MCKINLEY CO.: 6 mi. S. Thoreau (UMMZ 120297) ; 7 mi. S. Thoreau (CU G333) ; 
8mi.S.Thoreau(CU6232,6412);Ft. Wingate (USNM 14404, 53587); Valencia 
CO- 1 mi. N. Cebolleta (UMMZ 86605); 4 mi. WSW. Cebolleta (UMMZ 
86606); Ranger Station, Lobo Canyon, Mt. Taylor (UMMZ 86607); 1 1-5 mi- 
SSE. Grants (CU 6233); Grants (CU 5371, 5373, 5381, 5393, 5402; KUMNH 
19360-7*). 

Pseudacris triseriata triseriuta Wied x P. t. macuUita Agassiz 

Known from only seven published locaUties, the western chorus frog 
has been studied very little in New Mexico. Smith (1956) reassigned 
the subspecific name maculafa to P. nigrifa (=P. triseriata, Schwartz, 
1957, p. 11) from Colorado, Utah, and northward. With the exception 
of a smglc specimen of P. t. triseriata x P. t. macvlata from Colfax 



274 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

County, New Mexico, he left the Arizona-New Mexico population as 
P. t. triseriata. His criterion was the greater tibia length/body 
length ratio in frogs from the latter states. Five specimens from 
the Zunis have ratios of 38.8-43.5 (x 40.9 ±0.4), thus placing them as 
P. t. triseriata x P. t. maculata intergrades (cf. Smith, 1956, table 2). 

Recently transformed specimens, 13 and 14 mm. snout-vent, were 
collected July 13, 1949, in a meadow of the spruce-fir association at 
about 9200 feet. They were thought to be singing at the time of 
capture (Chenoweth, 1950, p. 532), but this seems improbable in view 
of their humaturity. Three males, also judged to be singing, were 
collected May 21, 1924, by Charles L. Camp at 7000 feet; they measure 
29, 30, and 31 mm. snout-vent and are reproductively mature. All 
Zuni specimens have three distinct dorsal stripes that are broken 
posteriorly in the three adults. 

Locality records : 

MCKINLEY CO.: Ft. Wingate (MVZ 9339-41); Valencia co.: Mark Elkin's 
Ranch, Mt. Taylor (CU 6338). 

Rana pipiens Schreber 

I have not attempted to assign Zuni leopard frogs to any particular 
geographic race in view of the notorious variability and inadequate 
definition of the various proposed subspecies (e.g., see Moore, 1944). 
In the study area these frogs may be many shades of green, brown, or 
gray; but they usually have 5-12 (x 9.1) large, regularly arranged 
dorsal spots and smooth sldn. With respect to the latter feature, 
they are quite different from the highly pustulose leopard frogs 
occurring near Snowflake, Navajo County (UMMZ 121949), and 13 
miles west of Chevalon, Coconino County (UMMZ 121951), Arizona, 
although from the same dramage system (Colorado River). The 
preserved specimens are small and may not represent the typical 
adult size range (table 4). A single female is 55 mm. snout-vent. 

Lack of a large series and the few locality records reflect only the 
amount of attention directed toward collecting this species. My 
field notes indicate that R. pipiens is one of the most abundant 
anurans in the Zunis; it was seen commonly around isolated pools 
and streams in the riparian association of Roughlands and Montane 
Life Belts. Coues (1875, p. 631) found it to be similarly abundant 
along the Zuni River and Rio Gallo near Ft. Wingate. I did not 
see R. pipiens in cattle tanks or roadside ditches, nor did I note any 
breeding activity; however, larvae of all sizes were found throughout 
the summer months, and a great concentration of recently transformed 
individuals, 9-25 (x 21.6) mm, snout-vent (Nl=5),was noted August 
21-23, 1959, at 7200 feet. 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 275 

Like Bujo woodhousei, leopard frogs also feed upon the tenebrionid 
beetle, Eleodes. 
Locality records: 

MCKINLEY CO.: 6 mi. S. Thoreau (CU 5773, 6306, 6340; CWGF) ; Thoreau 
(MCZ 32820-47,* 33300-05*); Ft. Wiugate (USNM 16760); Valencia co.: 
Grants (CU 5371-3). 

Class Reptilia: Suborder Sauria 

Crotaphylus collaris collaris Say x C. c. baileyi Stejneger 

The western collared lizard, originally described as C. baileyi by 
Stejneger (1890, p. 103) from the Painted Desert, Little Colorado 
River, Arizona, can be characterized by its two discrete interorbital 
scale rows, a tan or brown head with darkly spotted green to blue 
throat (in males), and a sharply contrasting, spotted body pattern 
(Fitch and Tanner, 1951, pp. 549-550). Bm-t (1928) s3monymized 
baileyi with nominate C. collaris and listed (Burt, 1933, p. 229) two 
localities in the Zuni region. Presentl}^ relegated to subspecific status, 
C. c. baileyi is the form usually found in the Zunis. This occurrence 
is not surprising because suitable habitats are more or less continuous 
between the type locality and the study area. 

According to Fitch and Tanner (1951, p. 549) and Stebbins (1954, 
p. 310), a broad zone of intergradation exists between C. c. collaris 
and C. c. baileyi. This is evident in the Zunis, for 3 of 18 specimens 
have one scale common to both interorbital rows. On the basis of 
this criterion and the fact that certain Arizona specimens show a 
similar condition (Burt, 1928, p. 11), the three Zuni individuals may 
be labeled C. c. collaris x C. c. baileyi. It seems likely that a relativelj^ 
smooth, east-west cline of variation is present in southwestern collared 
lizards (see discussion under "Zoogeography," p. 315). 

Reasons for retaining subspecies in C. collaris and the description of 
a new intermontane Utah-Colorado race, C. c. auriceps, were given 
by Fitch and Tanner (1951). Although C. c. auriceps occurs in Monte- 
zuma Comity, Colorado (Maslin, 1959, p. 26), and logically in San Juan 
County, New Mexico, its characters could not be discerned in the 
Zuni sample or in specimens from Chaco Canyon National Monument, 
San Juan County (UMMZ 122892). This is somewhat puzzling when 
one considers that other reptiles with similar distributions are present 
in both areas, e.g., Sceloporus undulatus elongatus. 

Fitch and Tanner (1951, fig. 1) show modal nmnbers of 14 for both 
supra- and infralabials of (7. c. baileyi, 13 and 12 for these respective 
counts in C.c. auriceps. They apparently plotted labials on both left 
and right sides of the head in the same histogram. The figm'e is 
labeled "Numbers of Specimens" but does not agree with the number 

738-287—65 3 



276 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

of speciiiieiis examined (cf. Fitch and Tanner, 1951, p. 556); thus, 
the relatively small difference in labial number appears twice as 
strongly emphasized. Actually, such a distinguishing feature easily 
could be missed in possible intergrades. 

Neither sexual dimorphism nor bilateral imbalance of labial scales 
was alluded to by the authors of G. c. auriceps. Etheridge (1960, 
p. 114) noted that males of C. c. collaris have larger heads and more 
robust lower jaws than females. In the Zunis, adult females are 
proportionately smaller than males (table 5) and possess fewer 
infralabials; seven have 11-14 (x 12.6), 12-16 (13.6) left and right 
infralabials respectively, and have 13-15 (14.0), 12-16 (14.3) left and 
right supralabials respectively. Eight males have 12-14 (13.1), 13-15 
(13.8), 14-16 (14.5), and 13-16 (15.0) left and right infra- and supra- 
labials. The bilateral imbalance is interesting in that the right side 
of the head seems to be favored, although this may not be meaningful 
in light of the sample size. 

Collared lizards are a typical reptile of boulder-strewn hillsides 
where vegetation is sparse. Such situations usually are found on the 
lower slopes of mesas and canyons and at the malpais edge in the 
Zuais. On several occasions, foraging individuals encountered on the 
tops of mesas seemed to be attracted by large numbers of grasshoppers 
feeding on the thicker vegetation. These lizards are not restricted to 
the Roughlands Life Belt, but on the plains they are confined to the 
vicinity of rocky arroyos. Sceloporus undulatus, Eumeces obsoletus, 
Masticophis taeniatus, Salvador a grahamiae, and C. collaris were 
collected in and around an arroyo at 6400 feet in a cholla-juniper 
dominated section of the Plains-Roughlands continuum. 

Females of C. collaris with the typical red breeding coloration 
were observed as late as August 11, 1959, but none collected in July or 
August held large eggs. Specimens taken by Emmet T. Hooper, 
May 7-9, 22, 1939, have only small ova. Two cross-banded hatchlings, 
40 and 41 mm. snout- vent, with evident yolk-sac scai's, were found 
August 25, 1951, at 6500 feet; thus, if about 51-74 days of mcubation 
are needed (Fitch, 1956a, p. 234), egg-laying may occm- in June or early 
July, ahead of the summer rains. 

Locality records: 

MCKINLEY CO.: Thoreau (MCZ 62420*); 5.5 mi. SE. Thoreau (CU 5748); 6 mi. 
S. Thoreau (CWGF); Ft. Wingate (USNM 16761); Gallup (USNM 27738); 14 mi. 
N. Gallup (UCM 6583-84); Valencia co.: 1.5 mi. S. Grants (UMMZ 86616); 8 
mi. SE. Grants (UMMZ 86615); 5 mi. E. Grants (CU 3051, 3376); 10.4 mi. S. 
Grants (CU 5747); 11.5 mi. SSE. Grants (UMMZ 122621-22); 7 mi. SE. Laguna 
(UMMZ 72649); 5 mi. NW. Rio Puerco (USNM 86937). 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



277 



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278 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

Holbrookia maculata campi Schmidt 

Subspccific assignment of H. maculata in the Zuni region is based on 
specimens examined by Ralph W. Axtell (in htt.), who has revived 
the name campi and restricted it to Colorado Plateau populations 
(Axtell, 1958). 

The lesser earless lizards that I collected are brown, red-brown, or 
gray dorsally, with 10-17 darker blotches or interconnected spots 
between extremes of limb insertions. They come from localities with 
soils of similar coloration. H. m. camjpi also approximates substrate 
color near Springerville, Apache County, Arizona, where specimens 
taken on soil-covered lava are dark brown or gray in general aspect. 
By contrast, an individual from the white sand along Choukai Wash, 
San Juan County, New Mexico (UMMZ 120277), is white to light 
gray, with darker gray dorsal spots (observations during three months 
in captivity). It is identical with H. m. ruthveni, the pallid "sub- 
species" of the White Sands, Otero County, New Mexico (see Smith, 
1943b; 1946, pi. 20A). Other pallid specimens of H. maculata come 
from the vicinities of Montoya (Quay County), Pedernal (Torrance 
County), and Carrizozo (Lincoln County), New Mexico. I suggest 
that H. m. ruthveni be relegated to the synonymy of H. m. approximans, 
the form inhabiting lowlands soutli of the Colorado Plateau, since 
this white sand population is not geographically unique in its only 
distinguishing character, coloration. 

In the Zunis, lesser earless lizards are inhabitants of the Plains Life 
Belt only. Four specimens come from the short-grass association in 
which snakeweed and rabbitbush is locally abundant. Another is 
from soil-covered malpais dominated by scattered junipers and apache 
plume. A third was in open grassland near a thicket of shadscale 
saltbush. Northeast of Prewitt where H. m. campi is associated 
marginally with Cnemidophorus velox and Sceloporus graciosus, a 
pureued earless lizard took refuge in the burrow of Dipodomys ordi. 
C. velox and S. graciosus frequent heavier cover such as saltbush or 
sage at this locality. 

Gravid females with the following snout-vent length, date of collec- 
tion, and egg number and size provide an estimate of local reproductive 
capacity in H. m. campi: 54 mm.. May 12, 1939, 6 avg. 9.8 x 5.2 mm.; 
57 mm., July 16, 1957, 3 avg. 13.4 x 7.4 mm.; 48 mm., July 16, 1957, 
3 avg. 12.8 x 7.5 mm.; 48 mm., July 16, 1957, 2 avg. 13.8 x 7.6 mm.; 41 
mm., July 5, 1959, 2 avg. 12.2 x 7.6 mm. It is possible that this 
iguanid deposits its eggs in response to summer rainfall (see discussion 
under Sceloporus undulatus). 

Locality records: 

MCKINLEY CO.: 5.5 mi. S. Thoreau (CU 5624); 4 mi. ENE. Prewitt (CU 5456j ; 
Upper Nutria (UMMZ 123135); Valencia co.: Grants (MCZ 6824*); Point of 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 279 

Malpais (UMMZ 8GG13); Acoma Pueblo (USNM 44696); Lava Beds (USNM 
44697); 8 mi. SE. Thoreau (UMMZ 122889). 

Comparative material examined: 

H. rn. carnpi. — New Mexico: san juan co.: 1 mi. N. Chaco Canyon Nat. 
Mon. (UMMZ 120277); Arizona: apache co.: 7 mi. N. Springerville (UMMZ 
121722). H. m. approximans. — New Mexico: otero co.: 3 mi. S. Alamagordo 
(UMMZ 64690, paratype of H. m. ruthveni); quay co.: 3 mi. S. Montoya (UMMZ 
69068-69); torrance co.: Pedernal (UMMZ 67850); Lincoln co.: Carrizozo 
(UMMZ 70080). 

Sceloporus undulatus tristichus Cope x S. u. elongatus Stejneger 

Some attention has been given to the occurrence of S. undulatus 
in northwestern New Mexico. The possibility that the widespread 
form S. u. tristichus might intergrade with the northern subspecies, 
S. u. elongatus, was suggested by Bm't (1933, p. 242). He considered 
S. u. consobrinus as the plateau lizard occurring in western McKinley 
County rather than S. u. tristichus, a name which he ascribed to 
intermediates between S. u. elongatus and S. u. consobrinus. Smith 
(1938, p. 16) defined S. u. tristichus in the sense employed here and 
outlined some intergrade populations in southern Utah and Colorado. 
He mapped S. u. elongatus as being present north of the Zunis but did 
not indicate specimens examined or published records from this part 
of the state. S. u. elongatus has been collected since at several locali- 
ties in San Juan County, New Mexico, and Apache County, Arizona, 
the closest to the study area being Chaco Canyon National Monument 
(CU 5618) and Canyon de Chelly National Monument (UMMZ 
120278). 

The Zuni sample is clearly intermediate between S. u. tristichus 
and S. u. elongatus (table 6) . An arithmetical measure of subspecific 
difference proposed by Ginsburg (1938) was revived to clarify this 
situation. With data drawn from Smith's study (1938, table 2), 
the foUowmg may be noted. For the best diagnostic character, 
i.e., nmnber of dorsal scales from occiput to rmnp, the smallest 
overlap between S. u. elongatus and S. u. tristichus is 10.6 percent. 
This value is 18.1 percent when S. u. elongatus and the Zuni series 
are compared and 51.6 percent when S. u. tristichus and the Zuni 
material are compared. As regards the next best character, number 
of scales around midbody, overlap (intergradation) values are 23.0, 
30.4, and 33.4 percent respectively. In both features Zuni specimens 
are nearer S. u. tristichus. 

Plateau lizards from the vicinities of San Rafael, Paxton Sprmgs, 
and El Morro National Monument, Valencia County, are most 
representative of S. u. tristichus, whUe those taken near Crownpoint, 
Prewitt, and Gallup, McKinley County, are closer to S. u. elongatus. 
Series collected around Thoreau, McKinley Comity, are intermediate 



280 



PROCEEDINGS OF THE NATIONAL MUSEUM 






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HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 281 

in most respects. This northwest-southeast cline may be extended 
west through the Defiance Plateau, Apache County, Arizona, where 
I also have taken S. u. tristichus x S. u. elongatus (UMMZ 120280), 
to Grand Canyon National Park, Coconino County, Arizona (Williams, 
1960, p. 26). 

In view of this cline, one might question the retention of S. u. 
tristichus and S. u. elongatus as distinct subspecies. Other salient 
ecological-morphological differences in New Mexico populations, 
however, prevent me from lumping these races. With the exception 
of a few nearl}'' unicolor specimens,* most S. u. elongatus from San 
Juan County, New Mexico, lack distinct dorsolateral and lateral 
fight stripes and the spotted lateral pattern figured by Smith (1946, 
pi. 53 E, G) for S. u. tristichus; furthermore, adults are larger 
(45-83, X 64 mm. snout-vent, 25 San Juan County specimens) and 
possesses more fourth-toe lamellae (22-25, x 23.3 ±0.29) than S. u. 
tristichus x S. u. elongatus (cf. table 5). They are decidedly more 
scansorial, frequenting large boulders and cliff situations in contrast 
to marked terrestrial propensities in typical individuals of S. u. 
tristichus and the Zuni population. 

Although Sceloporus undulatus was the most ubiquitous reptile 
studied, it did not enjoy the wide elevational range of Phrynosoma 
douglassi or Thamnophis elegans (fig. 3). The species was most 
abundant in Roughlands associations, especially the pinyon-juniper. 
While found to a limited extent at the upper edge of the Plains- 
Roughlands continuum, S. undulatus usually was outnumbered by 
S. graciosus, Crotaphytus coUaris, and Cnemidophorus velox in this 
habitat. Specimens were conspicuously absent from cliffs inhabited 
by Urosaurus ornatus. They were uncommon on large boulders and 
the lower trunks of trees. Most individuals utilized elevated sites 
such as fallen tree trunks for sunning purposes and as apparent vantage 
points but were distinctly terrestrial in foraging habits. Body 
temperatures of 10 individuals foraging at 7400 feet were 29-37, 
(x33.1±0.98)°C. 

Evidence presented by Anderson (1960, p. 66), Martin (1958, p. 57), 
and others indicates that reproductive activity in certain reptiles is 
correlated with the beginning of summer rains. Similar observations 
on S. undulatus have prompted the following remarks. Plateau 
lizards carrying well-developed eggs have been found as early as 
June 13, 1957. Most females over 40 mm. snout-vent held eggs when 
collected before mid-July, but none taken in August contained eggs. 

* A polychromatic variant occurs, usually uniform dark brown or gray but 
occasionally with slightly darker or lighter lateral stripes (e.g., see I.egler, 1960, 
p. 181; Smith, 1946, p. 217). It has been found in the Guadalupe Mounlaiiis, 
New Mexico (UMMZ 121745), but not in the Zuni region. 



282 PROCEEDINGS OF THE NATIONAL MUSEUM vol. u6 

It is possible that the onset of July rainfall provides these lizards and 
S. graciosus, Urosaurus ornatus, and Holbrookia maculata with 
optimum conditions for oviposition. Ten eggs of S. undulatus, 
averaging 12.5 x 8.5 mm., were recovered from a one-inch depression 
beneath a small sandstone slab July 23, 1956 at 7300 feet. The 
substrate was damp, loose sand, and the eggs were not buried. 

Large eggs taken from females often contained embryos with 
well-developed eyes as did all eggs from the clutch obtained July 23, 
1956. Similar predepositional embryonic growth was reported for 
S. undulatus in Arizona and S. orcutti (Stebbins, 1954, pp. 240-244). 
This might be expected if fertilized eggs are retained for an extended 
length of time as suggested here. The phenomenon may be adaptive 
in a semiarid region like the Zunis; i.e., to insure annual repro- 
duction within the limits of optimum activity temperatures, post- 
depositional development thus can be completed in a relatively short 
time when oviposition is keyed to adequate summer moisture con- 
ditions. Hatchlings, 23-28 mm. snout-vent, have been collected as 
early as July 26, August 9, and August 12 in different summers. 

Locality records: 

MCKINLEY CO.: 5.5 mi. S. Thoreau (CU 5623); 6 mi. S. Thoreau (CU 5088-9 
5448, 5622, 5681, 5697; UMMZ 121919, 122885-7); between Thoreau and 
Continental Divide (UMMZ 120311) ; Gallup (CU 5843) ; 14 mi. N. Gallup (UCM 
6630-37) ; Ft. Wingate (USNM 14406-7, 8250, 8491, 9548; UCM 6638) ; Manuelito 
(USNM 87031); 5 mi. SE. Baca (UIMHN 39685-6*); 4 mi. NNE. Prewitt 
(UMMZ 120284); 11 mi. W. Ramah (UMMZ 122884); Valencia cc: Cubero 
(USNM 87032-4); El Morro Nat. Mon. (KUMNH 15121-6*; UMMZ 122794); 
2 mi. W. El Morro Nat. Mon. (CU 5606) ; 2.5 mi. E. Laguna Pueblo (KUMNH 
28065*); 8 mi. SE. Thoreau (UMMZ 122890); Shuman's Ranch (UMMZ 86608); 
15 mi. E. Grants (KUMNH 15294-5*); 10.4 mi. S. Grants (CU 5608); 1.5 mi. 
S. Grants (UMMZ 86609); 4 mi. WSW. Grants (UMMZ 86610); 18 mi. SW. 
Grants (UMMZ 86611); 17 mi. SW. Grants fUMMZ 86612). 

Comparative material examined: 

S. u. elongatus. — New Mexico: san juan co.: Chaco Canyon Nat. Mon. 
(CU 5618; UMMZ 122896); 7 mi. E. Blanco (UCM 7337-47); 43 mi. SSE. 
Bloomfield (UCM 7350-55); 2 mi. S. Colo.-N. M. state line on U.S. Rt. 666 
(UCM 13294-13301); Arizona: apache co.: 10 mi. W. Window Rock (UMMZ 
120280, intergrades with S. u. trish'chus) ; Canyon de Chelly Nat. Mon. (.UMMZ 
120278, 120281). S. u. consobrinus. — New Mexico: eddy co.: 2 mi. NW. 
Carlsbad Caverns Nat. Park, headquarters area (UMMZ 121745). 

Sceloporus graciosus graciosus Baird and Girard 

Sagebrush lizards, represented by the nominate subspecies, usually 
are considered to be widespread in northwestern New Mexico (Smith, 
1946, p. 495; Stebbins, 1954, p. 317). Published records, however, 
are available only for Rio Arriba (Cope, 1900, p. 389; VanDenburgh, 
1924, p. 205) and Sandoval (Maslin and others, 1958, p. 335) counties. 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 283 

To these may now be added several localities in San Juan and Mc- 
Kinley counties (see below). It seems doubtful that S. g. graciosus is 
common south of the Colorado Plateau; specimens have yet to be 
found in Valencia County. Small, disjunct populations inhabit the 
sand dunes of Chaves County, New Mexico, and Ward and Winkler 
counties, Texas (Sabath, 1960). 

Many herpetologists have followed VanDenburgli (1922, p. 262) in 
separating the three subspecies of S. graciosus as follows: S. g. 
graciosus has 42-53 (x 47.8) dorsal scales from occiput to rump, 
S. g. gracilis has 52-68 (60.8), and S. g. vandenburghianus has 48-66 
(54.8). It is significant that the dorsal scale count is high in Zuni 
material (table 5). The same is true of San Juan County specimens, 
for 16 from the vicmity of Blanco (CU 5617, UCM 7274-79) have 
51-58 (x 54.3 ±0.57) dorsals and 10 taken south of Bloomfield (UCM 
7280-87, 7348-49) have 51-58 (54.9 ±0.86). Zuni and San Juan 
County series approximate S. g. vandenburghianus in number of 
dorsal scales, but the males resemble S. g. graciosus with their widely 
separated, bright-blue belly patches and distinctly striped dorsum. 

The specimens at hand represent variation that remains to be 
studied thoroughly. The entire polytypic complex is in need of 
revision as further evidenced by specimens from northern Arizona 
(Durham, 1956, p. 222) with equally high dorsal scale counts and by 
"nominate graciosus" from northeastern Utah (Legler, 1960, p. 180) 
with confluent belly patches. It is possible that Colorado Plateau 
specimens are subspecifically distinct from those of the Great Basin; 
hence, the present racial allocation is tentative. 

Since a pallid, arenicolous form of S. graciosus is known from 
southeastern New Mexico and adjacent Texas, it is of interest to re- 
port similar variation in northwestern New Mexico. Specimens 
collected on white sand along Choukai Wash, San Juan County 
(UMMZ 120283), approximate the description of aS'. g. graciosus pro- 
vided by Sabath (1960). The dark-brown dorsal stripes seen in 
Zuni specimens are lacking or are indicated only faintJy. These sage- 
brush lizards are light beige but retain the four white dorsal stripes. 
Two males have very light belly patches and throat mottling; the 
latter feature is apparently absent in southeastern New Mexico 
(Sabath, 1960). 

The terrestrial nature of S. g. graciosus is well documented (Eaton, 
1935, p. 12; Woodbury and Woodbury, 1945) and is particularly 
evident in the Zunis, where this lizard was observed in open, rela- 
tively level situations marked by the saltbush-sage association or 
pure stands of big sagebrush and loose clay or moderately sandy 
soil. In only one instance was S. g. graciosus abundantly associated 
with S. undulatus. At the base of the Vermilion Cliffs north of 
Prewitt, it was at least five times more common than its congener 



284 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

ill pjitcbes of spiny saltbush iii the Plains-Roughlauds ecotone. 
Cnemidophorus velox also was present but imcoininon. 

Females, 54 and 55 mm. snout-vent, collected June 10, 1957, each 
contained four eggs averaging 12.0 x 7.5 and 13.5 x 8.0 mm. respec- 
tively. Three others, 50, 52, and 57 mm., held, respectively, three 
(avg. 12.9 X 6.0), four (avg. 12.3 x 7.2), and four (avg. 12.0 x 6.0) 
eggs on July 9. Eight females taken July 23, 1959, had deposited their 
annual egg clutches, possibly in response to the onset of summer 
rainfall. No hatchlings were ever observed. 

Locality records: 

MCKINLEY CO.: 3 mi. N. Thoreau (CU 5699); 3 mi. NE. Thoreau (CWGF); 4 
mi. NNE. Prewitt (CU 5452, 5600, 5698; UMMZ 120282); Lizard Canyon, 
Satan Pass (ANSP 20987*); 14 mi. N. GaUup (UCM 6608-11); 5 mi. S. Manuelito 
(UMMZ 122888). 

Comparative material examined: 

Sceloporus g. graciosus. — New Mexico: san juan co.: 1 mi. N. Chaco Canyon 
Nat. Mon. (UMMZ 120283) ; Chaco Canyon Nat. Mon. (UMMZ 122902); Blanco 
Trading Post (CU 5617) ; 7 mi. E. Blanco (UCM 7274-79) ; 34-43 mi. SSE. Bloom- 
field (UCM 7280-87, 7348-49); 2 mi. S. Colo.-N. M. state line on U. S. Rt. 666 
(UCM 13292); 0.5 mi. S. Colo.-N. M. state line on U. S. Rt. 17 (UCM 1303-4). 

Uta stansburiana stansburiana Baird and Girard x U. s. stejnegeri 

Schmidt 

This is apparently another species with a north-south pattern of 
subspecific intergradation in the Zuni region; however, the present 
allocation must remain tentative until a series can be secured and 
the races of U. stansburiana are better characterized. Zuni speci- 
mens have 88 and 100 dorsal scales between the mterparietal plate 
and posterior edge of the thighs. One has contiguous prefrontals; the 
other has the prefrontals separated by two small scales (see Smith, 
1946, p. 277). 

Based on present subspecific concepts in U. stansburiana, this 
intergi'ade zone extends north into San Juan County, New Mexico. 
Five specimens from Chaco Canyon National Monument (CU 5619; 
UMMZ 122909) have 94-106 (x 99.2±1.5) dorsals; 6 from seven 
miles east of Blanco (UCM 7392-97) have 106-112 (108.7 ±0.67), 
and 10 taken two miles south of the New Mexico-Colorado state line 
(UCM 13377-86) have 100-114 (107.4 ±2.0). The large number of 
dorsals and prefrontal contact in only one San Juan County specimen 
indicate that influence from U. s. stejnegeri is relatively slight. 

The apparent rarity of Uta in the Zunis is difficult to explain. The 
two precise localities, at 6900 and 7100 feet, are relatively open grass- 
land with sandy soil, rabbitbush, snakeweed, and scattered one-seed 
junipers. The localities lie at the northwestern edge of mixed grass- 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 285 

land that extends along the Rio San Jose from the Rio Grande valley 
(see Castetter, 1956, fig.). This gi'ound uta is more abimdant in 
eastern Valencia county at lower elevations and is common in the 
San Juan Basin to the north. 
Locality records : 

MCKINLEY CO.: Thoreau (CWGF) ; 4 mi. ENE. Prewitt (CU, specimen lost); 
VALENCIA CO.: (KUMNH 12192*). 

Urosaurus ornatus linearis Baird x U. o. wrighti Schmidt 

Urosauriis has been the subject of considerable taxonomic debate. 
This genus was lumped with Uta by Schmidt (1953), Stebbins (1954), 
and Oliver (1943) subsequent to separation of the two genera by 
Mittleman (1942). Other authors such as Smith (1946), Smith and 
Taylor (1950a), and Stejneger and Barbour (1943) followed Mittleman. 
Although reasons for dividing Uta (in the sense of Mittleman, 1942) 
are equivocal as demonstrated by Savage (1958), arguments for this 
spht advanced by the latter author appear well founded. I consider 
Urosaurus as the proper generic name for forms occurring in the Zuni 
region and specifically allied with Uta ornata Baird and Girard (1852). 

The problem of discerning geographic variation in Urosaurus 
ornatus seems to result from Mittleman's failure (1942) to use signifi- 
cant quantitative methods in defining the various forms. This was 
demonstrated by Oliver (1943), whose method of treating U. ornatus 
was followed with modification by Langebartel and Smith (1954, pp. 
130-133). Murray (1953) questioned the use of degree of dorsal scale 
carination as a taxonomic character and, in addition to Duellman 
(1955, p. 8) and Gehlbach (1956, pp. 367-368), reported the failure of 
color to characterize subspecies in U. ornatus. Williams (1960, pp. 
27-28), experiencing difficulty with Mittleman's diagnoses, followed 
Langebartel and Smith (1954, pp. 130-133) in lumping U. o. linearis 
under U. o. schotti but recognized U. o. wrighti.^ 

Mittleman (1942, pp. 138, 146) recorded U. o. wrighti from McKinley 
and San Juan counties. New Mexico, and U. o. linearis from "generally 
south of Lat. 35° [in New Mexico] except for the southeastern quarter 
of the state." Within the Zuni region, U. ornatus exhibits varia- 
bility that, at first glance, suggests intergradation between U. o. 
linearis and U. o. wrighti (Gehlbach, 1956, p. 368); however, examina- 
tion of series of supposedly typical individuals reveals that most 
characters currently in use do not adequately separate these sub- 
species. Clinal variation is present in at least one important char- 

5 I provisionally consider linearis as a separate subspecies in view of the distinc- 
tiveness of specimens from south-central Arizona (fig. 8). A thorough examination 
of variation in U. ornatus from the Arizona- Mexico border region would clarify 
this situation but that is beyond the scope of my discussion. 



286 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

acter (fig. 8). Since U. oniatus is gi-eatly in need of a modern 
systematic revision, I hold subspecific allocation of Zuni specimens 
tentative at this time. 

Using Oliver's methods (1943, pp. 100-101), the following may be 
noted m U. ornatus from the Zunis. Enlarged dorsal scales are 
present in two distinct rows on each side of the smaller vertebrals in 
28 percent of a series of 29 specimens; they are in two rows with 
additional enlarged dorsals tending to form a third outer row in 66 
percent and in three quasi-distinct rows in 6 percent. Carination of 
these scales varies from a flat, weak keel to a sharp raised point with 
all types of intermediate forms observable. Similarly, complete 
gradation in the merging of dorsal tail-base scales with adjacent 
scutes is present. Thirty-one percent of the specimens have the 
enlarged dorsals separated by two rows of small vertebral scales, 
while 69 percent possess only one row of vertebrals. 



[j i gD 



16 



liil 



__ 17 

25 



B I J||li l iilllll l I 17 

19 



30 35 40 45 50 55 60 

DORSAL SCALES 

Figure 8. — Variation in the number of enlarged dorsal scales of Urosaurus ornatus from 
Arizona, New Mexico, and Utah. (Horizontal line = range; vertical line = mean; solid 
bar=two standard errors of the mean; hollow bar = one standard deviation. Sample 
size to right of range; provenance of sample to left: A, Pima and Santa Cruz Counties, 
Arizona; B, Cochise Co., Arizona; C, Yavapai Co., Arizona; D, McKinley and Valencia 
Counties, New Mexico; E, Kane Co., Utah; F, San Juan Co., Utah.) 

Thirteen individuals from the northeast of the Zuni Mountains 
have 40-53 (x 44.5 ±0.96) enlarged dorsals from the anterior insertion 
of the forelimbs to anterior insertions of hindlimbs; 16 from the 
southwest side have 35-47 (42.3 ±0.95). If these counts are com- 
pared with those of U. ornatus from Utah and Arizona, a north-south 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 287 

cline may be visualized (fig. 8). This gi-adient of increasing scale 
size extends into Sonora, Mexico (cf. Oliver, 1943, fig. 1). Two rows 
of enlarged dorsals on each side of the vertebrals are present in about 
SO percent of U. ornatus from southern Arizona. In approximately 
90 percent of those from northern Arizona and Utah, there is a partial 
or complete third, outer row. 

The rock uta is abundant on mesa edges and cliffs in the Rough- 
lands and lower Montane Life Belts. Its local distribution is 
determined apparently by the availabihty of large boulders or by 
areas of broken, exposed rock; hence, elevational limits are imposed 
only by the lack of such habitat. Each mesa partially isolated by 
level grassland is inhabited by a distinctive population marked by 
close resemblance to local rock color. While these utas usually were 
observed in a vertical position, they were found rarely in standmg 
trees and shrubs. Characterization of Urosaurus as a plant-dwelling 
genus (see Lowe, 1955b) does not hold for U. ornatus in the Zunis. 
This is well illustrated by its occurrence in montane canyon-head 
situations and absence from ridgetops. Arboreal vegetation is present 
in both places, but large rocks or outcrops are not prevalent m the 
latter. 

Data on reproduction indicates that the oviposition of U. ornatus 
may coincide with adequate summer moisture conditions. Gravid 
females with the following snout-vent length, date of coUection 
(1959), and egg number and size provide circumstantial evidence: 
54 mm., June 13, 5 avg. 6.7 mm. diameter; 47 mm., July 14, 3 avg. 
12.8 X 7.0 mm.; 45 mm., July 16, 4 avg. 10.6 x 5.9 mm.; 48 mm., 
July 16, 2 avg. 12.2 x 6.1 mm. Specunens taken July 23, 1959, had 
laid their eggs as had Sceloporus graciosus and >S'. undulatus at the same 
locality. Hatchlings, 25 and 26 mm. snout-vent, were found on 
August 10; three more, 24, 25, and 27 mm., were collected on August 
25, the following summer. 

Locality records : 

MCKINLEY CO.: 5.5 mi. SE Thoreau (CU 5625); 6 mi. S. Thoreau (CU 5129; 
UMMZ 120292) ; 7 mi. S. Thoreau (CU 5610, 5628) ; 8 mi. S. Thoreau (CU 5449) ; 
5 mi. NE. Thoreau (CU 5686) ; between Thoreau and Continental Divide (UMMZ 
120310); 4 mi. NNE. Prewitt (CU 5451, 5599; UMMZ 120203); 14 mi. N. Gallup 
(UCM 6092-93); Valencia cc: 8 mi. SE. Grants (UMMZ 86614); 17 mi. 
SW. Grants (CU 5127, 5695); 25 mi, S. Grants (MCZ 62308*); 2.5 mi. E. Laguna 
Pueblo (CU 5678); El Morro Nat. Mon. (CU 5605); 2 mi. W. El Morro Nat. 
Mon. (CU5445). 

Comparative Material Examined: 

U. 0. linearis. — Arizona: cochise co.: Carr Canyon, Huachuca Mts. (CU 
3539); 5 mi. SE. Sunnyside (CU 5126); Cave Creek Canyon, Chiricahua Mts. 
(CU 2469, 5122) , yavapai cc: 1 mi. N. Yarnell (CU 3036, 3513). U. o. schotii.— 
Arizona: pima co.: Arivaca (CU 1789); 5 mi. NE. Arivaca (CU 2461); santa 



288 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

CRUZ CO.: 3.5 mi. E. Patagonia (CU 5504); 9 mi. E. Sonoita (CU 877-8); Trench 
Mine Station (CU 5514). U. o. wrighli. — Utah: kane co.: near Kanab (UMMZ 
73380-84); san juan co.: near Moab (UMMZ 68573-78). 

Phrynosoma douglassi hernantlesi Girard 

The Zuni Mountains are the type locahty of Tapaya ornatissima 
Gu-ard (1858) by restriction (Reeve, 1952, p. 928). Presently known 
as Phrynosoma douglassi ornatissimum, this lizard and the closely 
related P. d. hernandesi have been reviewed most recently by Reeve 
(1952), who based the above locality restriction on Stejneger's desig- 
nation (1890, p. 113) of neotypes of P. d. ornatissimum. Critical 
examination of 29 adidt (table 5) and 17 subadult topotypes and the 
neotypes reveals that the Zuni population is composed actually of 
individuals with the characters of both P. d. ornatissimum and P. d. 
hernandesi (as defined by Girard, 1858, pp. 395-396; Reeve, 1952, 
pp. 924-928). 

With regard for diagnostic features, the adult specunens have: (1) 
temporal spines shorter than their basal diameter in 65 percent and 
about equal to basal diameter in 35 percent; (2) head width at jaw 
angle exceeding temporal width in 6 percent, about equal to temporal 
width in 44 percent, and less than equal in 50 percent; (3) dorsal 
dark spots light bordered mesially in 45 percent; and not light bor- 
dered in 55 percent. Only the first character favors allocation with 
P. d. ornatissimum; numbers two and three are more typical of P. d. 
hernandesi. These adults might be considered intergrades, but the 
added characterization of subadults and all specimens with special 
regard for locality data demonstrates that the above features vary 
ontogenetically as well as environmentally. 

Seventeen subadults, 24-45 mm. snout-vent, resemble P. d. 
hernandesi in color pattern as do five larger adults from the Montane 
Life Belt. Vivid dorsal coloration is virtually absent in these speci- 
mens; their dorsal blotches lack light mesial borders. In contrast, 
adults from Roughlands and Plains localities are bright orange or 
reddish brown in general aspect (brighter ventrally) and possess 
mesially and posteriorly light bordered blotches as in P. d. orna- 
tissimum. Tail length /snout- vent length is 0.369-0.471 (x 0.429) in 
subadults, 0.429-0.542 (0.470) in adults; hence, absolute tail length is 
unacceptable for comparative pm'poses as proposed by Reeve (1952, 
pp. 924, 928) unless ontogenetic variation is accounted for. Further- 
more, it appears that adults have relatively broader heads and longer 
temporal spines than subadults. None of the latter have the jaw 
angle width greater than the temporal width or have temporal spines 
equalmg their basal diameter in length. 

The idea that environmental gradients produce both P. d. hernan- 
desi and P. d. ornatissimum in a single geographic area is not new. 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 289 

Cope (1900, p. 417), concerned with P. d. ornatissimum, stated: 
"According to Stejneger this is the desert form of the species. In 
Arizona Dr. Merriam only found it on the Painted Desert, while he 
found the P. d. hernandesi in the cedar and pine belts in the San 
Francisco Moimtain region." Bailey (1913) included both races in 
his New Mexico, Upper Sonoran Life Zone, but only P. d. hernandesi 
in the Transition Zone. Eaton (1935, p. 16) commented on his 
inability to distinguish P. d. ornatissimum from P. d. hernandesi in 
northern Arizona, and Durham (1956, p. 222) described specimens of 
both "subspecies" from the Grand Canyon. Bragg and Dundee 
(1949, p. 57) and Lowe (1947) observed color variation correlated with 
ecological differences near Las Vegas and Socorro, New Mexico, 
respectively. In view of past and current difficulties in separating 
the two forms, I suggest that P. d. ornatissimum be placed in the 
synonomy of P. d. hernandesi, which has page priority (see Girard, 
1858, pp. 395-396). 

Short-horned lizards are certainly the most eurytopic reptiles of 
the Zuni region if not of New Mexico. They could be observed 
readily in aU major vegetational associations except the spruce-fir 
but were most abundant in the pinyon-juniper savanna. Specimens 
were taken in the alpine-tundra association at 11,300 feet on Mt. 
Taylor (La Mosca Lookout), a greater elevation than attained by any 
other species studied (fig. 3) . This is possibly the highest occurrence 
of any reptile or amphibian in the Southwest. 

The birth of ten young (24-26, x 24.8 mm. snout-vent) was observed 
in a clump of rabbitbush at 6400 feet, 11:00 a.m., August 14, 1957. 
AU were extruded within ten minutes and, with freedom from their 
investing membrane, immediately buried themselves in the loose, 
sandy soil. They were medium gray with only the dark-brown dorsal 
spots of the salient color pattern. Other dates (1957) of parturition 
and accompanying brood size are June 25 (15), July 11 (8), July 20 (6), 
August 3 (21), August 10 (11) for females taken between 7000 and 
7500 feet and maintained in outdoor cages at 7200 feet. A single 
juvenile, 20 mm. snout-vent, was collected on August 12 at 8200 feet. 
Specimens with ova, 0.6-1.5 mm. in diameter, were found throughout 
the summer months; one from 11,300 feet contained 11 eggs averaging 
4.2 mm. on July 17, 1956. 

Locality records : 

MCKINLEY CO.: 6 mi. S. Thoreau (CU 5119, 5233, 5447, 5632, 5703, 5709; 
UMMZ 120409) ; Ft. Wingate (USNM 14396, 14408) ; Prewitt (KUMNH 28063*) ; 
Rio Puorco, Continental Divide (AMNH 62853-4); Seven Lakes (CU 5620); 
Gallup (CU 5S41); 10 mi. N. Gallup (TNHC 11654-5); Valencia co.: Grants 
(MCZ 0791*; USNM 44805); 10.4 mi. S. Grants (CU 5601); La Mosca Lookout, 
Mt. Taylor (CU 5453); Mark Elkin's Ranch, Mt. Taylor (KUMNH 28062*); 



290 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Shuman's Ranch (UMMZ 86617); El Morro Nat. Mon. (KUMNH 11298-9*); 
Ojo Redondo (CU 5707); Mt. Sedgwick (USNM 44506); McCartey's (USNM 
16000-1); Lava Beds (USNM 44702). 

Cnemidophorus velox Springer 

Lowe and Zweifel (1952, p. 229) have characterized appropriately 
the systematics of whiptail lizards: "Perhaps there is no genus of 
lizard occurring in North America today that has been studied and 
continues to be studied with as much uncertainty and confusion as 
Cnemidophorus." In keeping with this statement, C. velox was, for 
many years, an unrecognized species. Probably it was reported in 
New Mexico by Yarrow (1875, p. 558) as C. octolineatus and by 
Hallowell (1854, p. 145), Cope (1900, p. 603), and VanDenburgh 
(1924, p. 213) as C. gularis. Burt (1931, p. 124) included it under C. 
sexlineatus perplexus and listed Zuni specimens from Grants and 
Gallup. Lowe (1955a) reviewed the nomenclatorial history of this 
Colorado Plateau species that Burger (1950a, p. 4) and others had 
confused with C. exsanguis, formerly known as C. sacki (Duellman 
and Zweifel, 1962). Maslm and others (1958, p. 342) reported C. 
velox from San Juan, Taos, Rio Arriba, and San Miguel Counties, 
New Mexico. Duellman and Zweifel (1962, fig. 10) utilized my 
distributional data on this species in their review of the "sexlineatus" 
group of Cnemidophorus. 

C. velox may be separated from C. exsanguis primarily on the basis 
of size and color pattern (pi. 3; table 7). Adults of the former are 
never spotted conspicuously, have more widely separated paraverte- 
bral light stripes and predominantly blue or blue-gray tails in con- 
trast to the spotted brown to greenish (distaUy) tailed adults of C. 
exsanguis. The two species are similar in scutellation (table 7). They 
are sympatric presumably in northern Arizona (Lowe, 1955a, p. 3), 
but this relationship is not certain in the Zunis. On two occasions a 
large seemingly spotted whiptail was seen foraging with typical 
adults of C. velox near Thoreau. Neither individual was collected, 
and other differences were not noted. Since some large C. velox 
develop light areas at the anterior ends of adjoining scales in the 
lateral dark stripes, these observations cannot be cited as indicating 
sympatry in the Zunis. 

Among the striped whiptails of New Mexico west of the Rio Grande, 
the scale counts of C. velox are closest to those of C. exsanguis and 
furthest from C. perplexus and C. inornatus; such features adequately 
separate C. velox from C. burti, a much larger spotted species (table 7). 
Unlike C. velox, C. perplexus is spotted as an adult and often has a 
wavy middorsal light stripe. C. velox and C. inornatus are the only 
unspotted species in western New Mexico. Both have blue tails, but 
C. inornatus is not larger than 67 mm. snout-vent, typically has a 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



291 



bluish venter, and rarel}^ (4 out of 37 New Mexico specimens) lacks 
the middorsal light stripe. In C. velox the venter is usually white, 
although occasionally tinged with blue, and the middorsal stripe is 
more often incomplete or absent (table 8). The condition of this 
stripe apparently remains constant during ontogeny (pi. 3A). 

Table 7. — Comparison of range and mean of meristic characters of striped ivhiptails 
(Genus Cnemidophorus) present in western New Mexico^ 



Species 


Dorsal scales 
around midbody 


Scales between 

paravertebral 

stripes 


Femoral pore^ 


Maximum 
snout-vent 
length mm. 


Spots 


C. burti 


98-115(104.7) 


5-11(8.0) 


32-44(38.3) 


130 


yes 


C. exsanguis 


62-86(74.8) 


2-8(4.6) 


30-44(36.8) 


101 


yes 


C. inornatus 


55-78(62.0) 


7-11(8.6) 


33-39(35.0) 


67 


no 


C. perplexus 


71-80(74.9) 


9-13(10.2) 


34-40(37.3) 


86 


yes 


C. velox 


67-80(72.5) 


5-10(7.9) 


28-38(32.9) 


84 


no 



• Data in part from Duellman and Zwelfel (1%2). 

There is relatively little geographic variation in C. velox (table 8). 
It is perhaps significant that populations from the immediate vicinity 
of the San Juan Basin are somewhat more uniform in structure and 
closer to the species mode than those to the south and west. Speci- 
mens from Catron County, New Mexico, have the fewest dorsal 
scales ^ and are smallest in size, closely approaching C. inornatus in 
these characters. Specimens from Kane County, Utah, have the 
largest number of dorsals and, including Navajo County, Ai'izona, 
and San Juan County, Utah, femoral pores. The four samples 
most peripheral to the San Juan Basin, those from the Zunis, Catron, 
Apache, and Kane Counties, show the greatest separation of para- 
vertebral light stripes and incidence of a complete middorsal stripe. 

Maslin (1959, p. 44) identified two specimens (UMMZ 76881) as 
syntypes of C. velox although he added that they were reidentified as 
C. stictogrammus by Richard G. Zweifel. Zweifel (in litt.) said that 
he had no notes on these specimens; he later examined and tentatively 
determmed them as C. sexlineatus. I concur with this identificatiou. 
Clearly the specimens in question are not syntypes of C. velox. Even 
their locality data, Lee's Ferry, Coconino County, Arizona, differs 
from that given by Springer (1928, p. 103) for the four original syn- 
types: "Three were taken at Oraibi, Arizona, and one at Pueblo 
Bonito, New Mexico." Some mixup is indicated since C. sexlineatus 
reaches its westernmost limits in San Miguel Count}'-, New Mexico, 
and cannot be expected at Lee's Ferry. 

6 I agree with Zweifel (1959, p. 66) that the number of dorsal scales around 
midbody is a more accurate count than the number from occiput to rump. This 
may be seen in the degree of variation in the Zuni population (table 5). 

73S-287 — 65—4 



292 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 293 

Of the 90 specimens of C. velox examined during tliis study, none 
were males. Comparative dissections revealed only famales of C. 
exsanguis and C. tessellatus, but bisexual samples of C. inornatus and 
C. tigris were noted. This problem is complicated further by the 
existence of two kmds of nonspotted whiptails in the Zuni region. 
One is typical of C. velox as previously characterized and has, in 
addition, abruptly enlarged scales preceding the gular fold. The 
other is represented by two males (UCM 6579-80) from 14 miles north 
of Gallup that resemble C. inornatus in having the bright-blue venter 
and smaller scales in front of the gular fold. These specimens, with 
68 and 75 dorsal scales around midbody, could be the males of C. 
velox, or they might represent C. inornatus in sympatry with C. velox; 
indeed, when more specimens become available and this perplexing 
situation is studied further, present species concepts involving C. 
velox and C. inornatus may be altered radically. 

In the Zunis, Colorado Plateau whiptails were most common in 
open areas of the Roughlands Life Belt especially where the saltbush- 
sage association occm-red in isolated patches in the pinyon-juniper 
savanna. They were associated occasionally with Sceloporus undu- 
latus in these habitats and with Sceloporus graciosus where scattered 
spiny saltbush and one-seed junipers dominated a sandy area at the 
lower edge of the Roughlands near Prewitt. Although seldom found 
above the Roughlands-Montane continuum, they sometimes occurred 
in ponderosa pine stands in open, logged-off spots or along dry creek 
beds to approximately 7800 feet. 

Gravid females with the following snout-vent length, date (1939) 
of collection, and egg number and size give some estimate of local 
reproductive capacity hi this whiptail: 77 mm.. May 28, 3 avg. 6.5 x 
15.2 mm.; 80 mm., June 13, 4 avg. 11.2 x 16.0 mm.; 73 mm., June 22, 
4 avg. 8.2 X 15.2 mm. A specimen collected June 30, 1956, had an 
enlarged oviduct but only small eggs measuring 0.5-1.5 mm. in 
diameter. None of the individuals taken in July held enlarged eggs; 
hence, C. velox apparently does not oviposit coinciden tally with sum- 
mer rainfall. A hatchling, 34 mm. snout-vent, was collected August 
13; another, 37 mm., on August 20, 1957. 

Locality records: 

MCKINLEY CO.: 4 mi. S. Thoreau (CU 5450); 5.5 mi. SE. Thoreau (CU 5614); 
6 mi. S. Thoreau (CU 5675; UMMZ 120410, 120289); 7 mi. S. Thoreau (CU 
5626); 4 mi. NNE. Prewitt (UMMZ 120286); between Thoreau and Continental 
Divide (UMMZ 120309); Ft. Wingate (USNM 16762-63, 80357); 16 mi. S. 
Gallup (CU 5609); 10 mi. W. Gallup (MVZ 65805); 14 mi. N. Gallup (UCM 
6578-80); Gallup (UMMZ 120287); Crownpoint (UNMCV 1162); Valencia 
CO.: 8 mi. SE. Paxton (UMMZ 86619); 1 mi. N. Cebolleta (UMMZ 86620) ; 4 mi. 
W. McCartey's (UMMZ 86618); 11.5 mi. SSE. Grants (CU 5458). 



294 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Comparative material examined: 

C. velox. — Arizona: apache co.: Little Colorado River (UMMZ 85054); 
NAVAJO CO.: 1 mi. N. Winslow (UMMZ 74025); New Mexico: cateon co.: 3 
mi. NE, Horse Springs (UCM 6095-6102); rio arkiba co.: 7 mi. N. El Rito 
(UMMZ 69081); san juan co.: 7 mi. E. Blanco (UCM 7250-59); 43 mi. SE. 
Bloomfield (UCM 7260-62); 1 mi. N. Chaco Canyon Nat. Mon. (UMMZ 120288); 
Chaco Canyon Nat. Mon. (UMMZ 122915); Utah: kaxe co.: 6-7 mi. NW. 
Kanab (UMMZ 73323-24); san juan co.: 3 mi. W. Monticello (UMMZ 121484- 
85); Natural Bridges Nat. Mon. (UMMZ 120285). 

Eumeces multivirgatus multivirgatus Hallowell x E. in. epipleurotus 

Cope 

The taxonomic status of southwestern members of the E. multi- 
virgatus complex recently has been discussed by Lowe (19o5c), Maslin 
(1957), Mecham (1957), Heyl and Smith (1957), and Tanner (1957). 
These authors are in essential agreement that E. gaigeae (=£'. m. 
epipleurotus; see Axtell, 1961) and E. taylori are pattern variations of 
E. multivirgatus, but then- opinions diverge as to the proper allocation 
of the three available names. Mecham (1957, pp. 113-116) proved 
that E. taylori is simply an allelomorphic pattern variant of E. multi- 
virgatus; thus, there is no reason, in the interest of biosystematics, for 
retaining E. m. taylori as a nongeographic subspecies as Lowe (1955c) 
and Tanner (1957 p. 116) do. Maslin (1957, p. 87) and Mecham 
(1957, p. 112) suggested that E. gaigeae represents a stage in the onto- 
genetic pattern change of E. multivirgatus. Such loss of color 
pattern is well documented in the E.fasciatus group but has not been 
demonstrated effectively in the many-lined sldnks. 

A series of 25 adults and 5 juveniles of E. m. multivirgatus from the 
Zuni region corroborates previous suggestions concerning the deriva- 
tion of the color pattern of E. m. epipleurotus. Complete pattern 
reduction can be observed in this variable sample and is comparable 
to Tanner's diagrams B through D (1957, fig. 1). An adult female, 
collected August 12, 1960, lost the mid-dorsal light stripe in 21 months 
of captivity. 

To test possible correlation of pattern change with increasing size, 
and presumably age, the hybrid index method of Sibley (1950, p. 112) 
and others was adapted for present use. The characters examined 
were middorsal light line, dorsal dark line on the first and/or second 
scale rows, lateral hght line on the sixth and/or seventh scale rows, 
and tail color pattern. The degree of intensity of each stripe was 
graded per specimen on a scale of (= present, as in typical E. m. 
multivirgatus) to 3 (—absent, as in typical E. m. epipleurotus). Tail 
color-pattern change (from blue to blue gray mth lines, to brown 
with lines, and ultimately to brown with lines and spots) was gi-aded 
in like manner. By adding the character values for a specimen, a 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



295 



total of indicated E. m. multivirgatus, 12 denoted E. m. epipleurotus , 
and 1-11 indicated ascending degrees of intermediacy. The specimen 
totals plotted against snout-vent length show ontogenetic change of 
color pattern from E. m. multivirgatus to E. m. epipleurotus with 
increasing size u-respective of sex (fig. 9). 



12 r 



10 



O 

8-* 



o 
o o 



o 
A A A 

A A O 

A A A 

% o o 



A 
A • 



Ai 



e a I 

I 1 



20 



30 



40 50 

SNOUT- VENT LENGTH mm. 



60 



70 



Figure 9. — Relationship of color-pattern to snout-vent length In Eumeces muUivirgaius 
from the Zuni region. (The color-pattern of nominate multivirgatus is 0, that of epi- 
pleurotus is 12.) 



A Males 

# Females |With blue tails 

H HatchlingsJ 



A Males 
O Females 



Without blue tails 



Suggested bases for retaining E. m. epipleurotus as a southern race 
of E. multivirgatus are: (1) ontogenetic pattern reduction (not 
present in nominate subspecies; Maslin, 1957; Mecham, 1957); (2) 
distinctive juvenile pattern (see Maslin, 1957, fig. IE); (3) posterior 
border of last supralabial confluent with posterior border of last infra- 
labial (supralabial projecting caudad at least 15 percent of its length 
in nominate subspecies; Maslin, 1957, p. 89). The first and second 
criteria apply to the Zuni skinks; however, for the thu-d, only 38 
percent of the adults possess the labial feature of E. m. epipleurotus on 
both sides of the head and 25 percent resemble E. m. multivirgatus, 
with the remainder being asymmetrical. While this third criterion 
remains to be tested in other New Mexico populations, some influence 
from the northern race is expected if physiographic and ecologic 



296 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ua 

factors operating in this situation are similar to those effecting north- 
south intergradation in other Zuni reptiles. 

Polychromatism in E. multivirgatus presents an interesting possi- 
bility for increased adaptation. It would be of distinct advantage 
for this skink, a wide-ranging form, to possess a variety of phenotypes 
each of which is superior in adaptive value to others in a given habitat 
occurring regularly in the range. Prevalence of the ^'taylori" variant 
in lower, more xeric situations in southern New Mexico is significant 
in this regard as is the apparent difference in proportion of variants 
in local populations (Gehlbach, in ms.; Mecham, 1957). The tend- 
ency for darkening in southern portions of the range is carried out 
by ontogenetic pattern reduction as well as by the occurrence of 
"taylori." 

Many-lined skinks occurred in a wide variety of Roughlands situa- 
tions, but most often they were taken in heavy cover such as the 
oak-mahogany and riparian associations where broad-leaf litter was 
abundant. A few were found under rocks and logs in the dryer, 
pinyon-juniper and ponderosa pine associations. The species was 
decidedly less saxicolous than Eumeces obsoletus and apparently did 
not occur in ecological sympatry with this larger skink. Absence of 
E. multivirgatus from the Plains Life Belt demonstrates this local 
relationship. Although collected throughout the day, many-lined 
skinks were seen more often in the early morning hours. 

On June 28, 1954, a female brooding three eggs was found in a 
shallow, cup-shaped depression under a loose rock in the dry bed of 
Cottonwood Creek at 7200 feet. The soil was damp only around the 
eggs, which measured 14.0 x 9.5, 14.5 x 8.5, and 15.0 x 7.8 mm. Five 
eggs averaging 14.2 x 8.6 mm. were dug from a Pueblo Indian ruin at 
a depth of approximately five inches on August 7, 1957. These were 
accompanied by a many-lined skink that escaped capture in the maze 
of rocks and loose, moist soU. Another female of the "epipleurotus" 
type and four recent hatchlings, 22 to 27 mm. snout-vent, were taken 
together on August 17. These hatchlings and one other, 29 mm., 
collected July 29, have the sharply contrasting juvenile pattern of 
E. m. epipleurotus. 

Locality records: 

MCKINLEY CO.: 6 mi. S. Thoreau (CU 5435, 5596, 5615, 5693; UMMZ 123529); 
8 mi. S. Thoreau (CU 5145); Ft. Wingate (USNM 16015-17); 10 mi. N. Ramah 
(UMMZ 122883); Valencia co.: 8 mi. SE. Thoreau (UMMZ 120408); 8 mi. NE. 
Grants (CU 5436); 17 mi. SW. Grants (CU 5607); 1.5 mi. S. Lookout Mt. (CU 
5691); 3 mi. SW. Page (CU 5692); 2 mi. W. EI Morro Nat. Mon. (CWGF). 

Emueces obsoletus Baird and Girard 

Burt (1929) and Taylor (1935, pp. 205-220) have reviewed geo- 
graphic variation in this species. Adult specimens collected in the 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 297 

Zuiiis fit tlieii" dcscriptiuns of New Mexico and Arizona material; 
however, two hatchliugs differ slightly from published accounts of 
juvenile coloration. They are 35 and 36 mm. snout-vent, with a 
glossy-black ground color that is equally as intense on the venter 
as on the dorsum. White spots on the head are confined to the upper 
labials, outside edges of the parietals, prefrontals, and supranasals, 
the six to eight scales immediately craniad and caudad of the ear, 
and the fu'st four infralabials. 

The Great Plains skink apparently reaches the northwestern limit 
of its New Mexico range in Valencia County and probably does not 
occm* on the Colorado Plateau further to the north. Locally it 
inhabits rocky situations in the Plains-Roughlands continuum and 
especially the cholla-juniper association of the Plains Life Belt. 
It was collected among rock outcrops in arroyos and along the malpais 
edge where there was relatively little shrub cover. In contrast to 
the plains-dwelling iguanid, Holbrookia inaculata, it seems to shun 
open areas. 

A female, accompanied by three recently hatched young, was 
discovered July IS, 1956, near the malpais edge on scoriaceous lava. 
Upon pursuit, she took refuge under a lava chunk, which, when 
overtm'ned, revealed her in the process of consuming one of the 
hatchliugs. The other two hatchliugs subsequently were collected 
and are the ones described above; the adult escaped. 

Locality records: 

VALENCIA CO.: Grants (USNM 44806); 1.5 mi. S. Grants (UMMZ 8G621); 
11.5 mi. SSE. Grants (CU 5455); 8 mi. SE. Paxton (UMMZ 86623); 4 mi. W. 
McCartey's (UMMZ 86622). 

Suborder Serpentes 

Thamnophis dorsalis dorsalis Baird and Girard 

I foUow Fitch and Milstead (1961) in replacing Thamnophis 
cyrtopsis Kennicott with the older Baird and Girard name. This 
arrangement is not enthely convincing because the black-necked 
gartersnakes, T. dorsalis, T. eques, and T. marcianus, of the Southwest 
and Mexico have been confused widely (see Wright and Wright, 
1957, p. 770); Fitch and Milstead (1961) admit that the original 
description of T. dorsalis is not sufficiently diagnostic to identify the 
missing type with certainty. 

The single male of T. d. dorsalis, collected during the present study, 
was found while I was investigating a breeding aggregation of Scaphiopus 
hammondi and Bvfo punctaius on the eastern edge of the Grants 
malpais at 6400 feet (Gehlbach, 1956, p. 369). This locaUty is at 
least 10 miles from the nearest stream of appreciable size, the Rio 
San Jose. The specimen has 174 ventral scales, 76 subcaudals, and 
19-19-17 dorsal scale rows. It is unlike most T. d. dorsalis in that it 



298 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

shows no dorsal spotting caudad of the neck region and has a very low 
subcaudal count (see Milstead, 1953). Reduction of posteriodorsal 
dark spots has been observed in specimens from Farmington, San 
Juan County (UMMZ 6G973-4), and Pinos Altos, Grant County 
(UMMZ 75626), New Mexico, but this reduction was not of the 
magnitude seen in the Zuni snake. 

T. dorsalis is not as abundant as T. elegans in northwestern New 
Mexico. It does not occur frequently in the pinyon- juniper associa- 
tion and, therefore, is separated somewhat ecologically from T. 
elegans. In addition to the Farmington specimens, others from 
the Acoma Indian Reservation (UNMCV, specimen lost), Suwanee, 
Valencia County (UNMCV 44, 117), Aztec (AMNH 5298), and junc- 
tion of the Los Pinos and San Juan rivers, San Juan County (A. Harris, 
in litt.), were taken below 6000 feet. Milstead's record (1953, p. 372) 
from ten miles north of El Rita [sic], Rio Arriba County, is questionable; 
I consider the specimens (UMMZ 84330) to be juveniles of T. elegans. 

Locality records: 

VALENCIA CO., 11.5 mi. SSE. Grants (CU 5053). 

Thamnophis elegans vagrans Baird and Girard 

Of all snakes represented in the Zuni region by a single geographic 
race, T. elegans exhibits the greatest morphological variation and 
widest range both geographically and altitudinally (fig. 3). Variation 
is especially apparent in the number of ventral and subcaudal scutes 
of both sexes (table 9). The means of these features approximate 
those given by Fitch (1940, figs. 3, 4), but the ranges are greater 
than his combined Arizona-New Mexico sample. Six specimens have 
23 anterior scale rows and one has divided preoculars, traits which 
Fitch (1940) considered rare in T. e. vagrans. 

Color pattern is subject to extremes of manifestation in the Zuni 
population. Certain individuals are nearly uniform olive brown 
dorsally with only a trace of the middorsal liglit stripe and dark dorsal 
spots. Others are marked with brilliant lemon-yellow stripes and 
clear black spots. In these the dark postparietal crescentic marks 
are strikingly similar to those of T. dorsalis and may have been part of 
the reason for misidentification in other studies. Black spots along 
the belly midline are present in varying degrees. Although a great 
variety of colors and patterns can be observed, no specimen possesses 
the ash-gray ground color seen by me in T. e. vagrans in southern 
Utah; all are some shade of brown. 

Juveniles usually are spotted distinctly, with the dorsal and lateral 
light stripes reduced in intensity and the black nuchal crescents 
clearly indicated. To my eye, juveniles are colored more cryptically 
than adults, which are generally less spotted. There is apparently 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



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300 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

some ontogenetic change in the color pattern of this subspecies, but 
the variety of polychromatism renders its interpretation especially 
difficult. 

Twenty young born on August 26, 1957, to a female, 1051 mm. in 
total length, are 142-160 (x 152) mm. snout-vent, 49-58 (55) mm. 
tail length. It is noteworthy that the above female is the largest 
T. e. vagrans on record; her brood averages considerably larger than 
those reported by Wright and Wright (1957, p. 799). Other dates of 
birth and brood size are July 11 (13) and July 2 (15) for females 
measuring 744 and 979 mm. total length, respectively. All brood 
records are from specimens captured at 7200 feet in the riparian associ- 
ation bordered by oak-mahogany and pinyon-juniper associations. 
The adults were maintained in outside cages at the same elevation. 

As indicated by Coues (1875, pp. 614-615) in observations made 
along the Zuni River, the wandering garter snake is typically semi- 
aquatic in the Zunis. Specimens of all sizes were encountered fre- 
quently in riparian situations where the amount of exposure varied 
according to elevation. The form was most abundant in Montane 
and Roiighlands Life Belts, but a few individuals were found on the 
plains in or near arroyos stemming from the uplands. Of 23 specimens 
with food in their stomachs, 17 had eaten Rana pipiens, both adults 
and larvae; 6 contained Sceloporus undulatus and Biifo woodhousei; 
2 held Peromyscus species; 1 held Perognathus flavus; and 1 each had 
eaten Phrynosoma douglassi and Ambystoma tigrinum. On several 
occasions, this garter snake was seen chasing anuran larvae under 
water. 

Locality records: 

MCKINLEY CO.: Thoreau (MCZ 62258-64, 62454-70*); 6 mi. S. Thoreau (CU 
5054, 5444, 5586, 5616, 5668) ; Ft. Wingate (USNM 16764-5, 14397-14401); 15 mi. 
N. Gallup (MVZ 63704*); 5 mi. SW. Ramah (UMMZ 123132) ; Valencia co.: 1.5 
mi. SW. San Mateo (UMMZ 86626-7); 4 mi. W. McCartey'g (UMMZ 86628); 
4 mi. WSW. CeboUeta (UMMZ 86624-25); Paguate (UMMZ 86629); Canyon 
Lobo Ranger Station, Mt. Taylor (UMMZ 86630); 7 mi. W. San Fidel (AMNH 
74501, 74786, 75966) ; El Morro Nat. Mon. (KUMNH 8557*) , 7.8 mi. SE. Grants 
(UMMZ 123131). 

Diadophis pimctatus regalis Baird and Girard 

Confusion in applying the name regalis has resulted apparently 
from a scarcity of museum specimens. Blanchard (1942, pp. 56, 62) 
considered D. regalis regalis as distinct from D. regalis laetus by 
absence or great reduction of the neck ring. Only two New Mexico 
specimens were available to him, however. Wright and Wright 
(1957, p. 161) followed Blanchard, although Schmidt and Smith 
(1944, pp. 89-90) previously had regarded presence or absence of the 
neck ring as anomalous and size and ventral scale counts as more 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 301 

significant. These latter authors synonymizod D. r. laetas with 
D. r. regalis but named D. regalis blanchardi. They were followed by 
Stebbins (1954, p. 489), who further suggested, as had Garman 
(1883, p. 73), that perhaps all ringneck snakes constitute a single 
species. 

Another rearrangement was fostered by Smith and Taylor (1945, 
p. 48), who maintamed Blanchard's views on D. regalis and D. r. 
laetus in "view of the paucity of data on these snakes" but synony- 
mized D. r. blanchardi with nominate D. regalis. Brown (1950, 
p. 145) supported their position. The form D. dugesi had been 
linked subspecifically with D. regalis (Taylor and Smith, 1938, p. 240) 
and D. punctatus (Villada, 1878) but was considered as a distinct 
species by Blanchard (1942, pp. 51-54). Utilizing new material from 
western Texas and southeastern New Mexico, Mecham (1956) com- 
bined D. regalis with D. punctatus but left subspecific boundaries in 
question. 

Taxonomic appraisal of Zuni ringneck snakes and others from New 
Mexico necessitated the following brief review of variation in south- 
western and Mexican Diadophis. I have been fortunate in seeing 
more material than was available to previous workers. Series from 
single localities, nevertheless, are nonexistant; hence, certain mor- 
phologically similar specimens were grouped politically (Aiizona, 
Utah, Chihuahua). Fm-ther evidence for considering the montane 
forms D. regalis and D. dugesi as subspecies of D. punctatus has been 
found. The racial boundaries left open by Mecham (1956) can now 
be approximated, and D. r. laetus, along with D. r. blanchardi, may 
be placed in the synonymy of D. p. regalis. 

As suggested by Cope (IGOO, pp. 544-545), the neck ring is a poly- 
chromatic character with dominance in some areas and variable 
manifestation in others. It occurs in some individuals from Sandoval 
County (New Mexico), Yavapai County (Arizona), and Chihuahua 
(Mexico), but is absent in others. Zuni specimens have neck rings 
(pi. 4) as do those from Catron, Sierra, and Grant Counties in south- 
western New Mexico. Material examined from central and south- 
eastern New Mexico usually lacks the ring, but one individual from 
Eddy County has it interrupted middorsally (Mecham, 1956). 
Incomplete neck rings also occur in the prairie race, D. p. arnyi (=i5. 
p. docilis, Mecham, 1956), and specimens from Utah, Trans-Pecos, 
Texas (Blanchard, 1942, pp. 63, 73; Mecham, 1956), and Jalisco, 
Mexico (UIMNH 47866). Most specimens from Utah and Trans- 
Pecos, Texas, lack the ring, whereas most Arizona and Mexican indi- 
viduals have it well developed. 

Similarly, extent of the yellow color of the venter on the dorsal 
scale rows varies from a single spot on a few anterior scales of the first 



302 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

row to nearly complete coverage of the first or first and second rows ; 
for example, one of the three Zuni specimens has yellow limited to 
a spot on each scale of the first dorsal row, while the others have the 
first row nearly covered with, and the second row spotted with, 
yellow (pi. 4). Most Arizona and New Mexico ringnecks show the 
former condition; most Utah snakes, the latter. In D. p. dugesi the 
yellow pigment is confined usually to the ventrals, but specimens 
from Hidalgo (UIMNH 17635), Jalisco (AMNH 3711-12), and 
Nayarit (AMNH 19724) resemble anteriorly those from Arizona-New 
Mexico. An individual from Durango, Mexico, (ISM, uncataloged) 
has faint yellow spots on the first dorsal row and thus appears to be 
intermediate between D. p. dugesi and D. p. regalis. A similar speci- 
men from Chihuahua (BYU 14243) has yellow confined to the venter 
caudad of the neck region. 

The Guadalupe Mountains population (Eddy Co., N. M. -Culberson 
Co., Tex.) also exhibits intermediacy in coloration as well as certain 
meristic featm'es described below. One specimen from Walnut 
Canyon (CCNP 2197) has yellow spots on all scales of the fu'st dorsal 
row, thus resembling most Arizona-New Mexico ringnecks. Another 
(CCNP 2201) has such spots only on the anterior quarter of the body, 
while a third has a uniformly gray dorsum. The McKittrick Canyon 
snakes have no yellow pigment dorsally except in the region of the 
partial neck ring, where it reaches approximately the fifth scale row 
on each side. Restriction of yellow pigmentation to the ventral 
scales is supposedly typical of D. p. arnyi (Stebbins, 1954, pp. 360, 
489) as well as D. p. dugesi (Blanchard, 1942, p. 51). 

In southwestern and Mexican Diadophis, the number of labial 
scales (usually 7/8), loreals (1-1), and pre- and postoculars (2-2) is 
geogi'aphicaUy stable, although often asymmetrical. By contrast, 
body length and three of fom* other meristic characters vary clinally 
(table 10). Two steps in the temporal and dorsal scale row clines are 
important because they fall at major breaks in the range of nonclinal 
ventral scale variation (fig, 10). One of these steps occurs in the 
Guadalupe Mountains south through the Sierra Vie] a and Chisos 
Mountains, Trans-Pecos, Texas, between the ranges of D. p. arnyi in 
west Texas and D. p. regalis in central New Mexico. The other lies 
between the ranges of D. p. regalis in Chihuahua, Alexico, and D. p. 
dugesi in southern Mexico. The steps are indicated by changes in 
frequency involving 15-15 and 17-17 dorsal scale rows and 1-1 
temporals. 

That certain meristic characters exhibit clines in different directions 
is evident. Ringneck snakes from the Sierra Vieja-Chisos region are 
closest to D. p. arnyi in dorsal and temporal counts but resemble D. 
p. regalis in number of ventrals. Almost the opposite situation exists 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 



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304 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

in the Guadalupes, where most specimens approximate D. p. arnyi in 
dorsal and ventral counts and D. p. regalis in frequency of temporal 
combinations; however, a male from Walnut Canyon is closest to 
D. p. regalis in having 206 ventrals. On the basis of clinal variation 
and intermediate coloration, ringneck snakes from the Guadalupe 



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160 170 180 190 200 210 220 230 240 250 

VENTRAL SCALES 

Figure 10. — Variation in the number of ventral scales of Diadophis punctatus from the 
Southwest and Mexico. (Solid circles = intergrades; hollow circle = a specimen from 
Durango. See table 10 for other provenance data and figure 8 for explanation of diagram.) 

Mountains south through Trans-Pecos, Texas, are considered inter- 
grades between D. p. arnyi and D. p. regalis. Perhaps this zone of 
intergradation extends further north. Bragg and Dundee (1949) 
reported D. p. arnyi seven miles south of Las Vegas, San Miguel 
County, New Mexico; typical D. p. regalis occurs 65 miles to the west 
in Sandoval County. 

West of the Guadalupe Mountains, ventral scale counts are uni- 
formly high among specimens from central and western New Mexico, 
Arizona, Utah, and northern Chihuahua (fig. 10). Zuni ringnecks 



HERPETOLOGY OF ZITNI MOUNTAINS — GEHLBACH 305 

are thus allocated with D. p. regalis, which is best distinguished by the 
large, sexually dimorphic number of ventrals. Thirty- three females 
have 220-247 (x 233.9±1.3) ventrals, while 28 males have 204-229 
(217. 3± 1.2). Over a range of 43 ventrals there is only a nine-scale 
overlap between the sexes. On the other hand, 25 males have more 
subcaudals (62-81, x 70.9 ±0.98) than 28 females (55-73, 64.8 ±0.73). 
Here the overlap, 11 scales in a range of 26, is relatively greater, 
rendering number of subcaudals inferior to number of ventrals as an 
external criterion of sex. Equally striking sexual dimorphism obtains 
in D. p. arnyi and D. p. dugesi, which have fewer ventrals than D. p. 
regalis (fig. 10). 

A single ringneck from southern Chihuahua (BYU 14243) has only 
196 ventrals, suggesting intergradation with D. p. dugesi (Tanner and 
Robison, 1960, p. 68). It further resembles D. p. dugesi in number of 
dorsal scale rows (17-17) and the coloration described above. A 
specimen from Durango is likewdse intermediate in ventral count 
and color (see above and fig. 10) but is closest to D. p. regalis horn 
Chihuahua in number of dorsals (17-15) and temporals (1-1). Un- 
fortunately, the other Durango specimen (UMMZ 102527) is in such 
poor condition that exact analysis of its scutellation is impossible. 
The Durango ringnecks and the above-mentioned individual from 
Chihuahua are considered intergrades since they bridge the previous 
morphologic and geographic gap between D. p. regalis in Chihuahua 
and D. p. dugesi in Nayarit. 

Martin and Harrell (1957, p. 469) listed D. punctatus among 
vertebrates that may have dispersed between the eastern United 
States and Mexico. The relationship of D. p. dugesi to D. p. arnyi 
through intergradation with D. p. regalis implies that dispersal via 
the Sierra Madre Occidental and the montane Southwest deserves 
equal consideration. Lower ventral and subcandal scale counts in 
D. p. dugesi and D. p. arnyi could be considered convergence in view 
of the distributional hiatus in northeastern Mexico. This hiatus 
may not be real, however, because a specimen described by Blanchard 
(1942, p. 59) from San Luis Potosi, Mexico, appears to be somewhat 
intermediate between D. p. arnyi and D. p. regalis and indicates that 
other such individuals occur in the region and possibly to the north 
also. Blanchard (1942, pp. 129-132) described breaks in range, 
considered by him important in maintaining D. p. regalis and D. p. 
dugesi as distinct species, that recently have been filled in. 

The secretive nature of Diadophis can be cited as a reason for its 
seeming rarity in the Southwest and Mexico. In the Zunis, a male 
collected August 17, 1959, was crawling over sandstone rubble near 
a small stream at 9:45 a.m. Although the locality, in pinyon- 
juniper-ponderosa ecotone at 7200 feet, had been worked intensely 



306 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

in preceding summers, ringneck snakes previously had not been found 
there. Another individual found in the riparian association displayed 
typical tail-curling behavior as did the above specimen (pi. 4). A 
female found September 13, 1958, at 6400 feet in an area of broken 
lava and sandstone contains 18 eggs, the largest being 2.5 nun. in 
diameter. It is 650 mm. in total length and has three divided ven- 
trals, including the first one anterior to the divided anal plate. 

Three live specimens from Cottonwood Gulch (6 mi. S. Thoreau) 
suggests that some color change may occur during the growth of D. p. 
regalis. The smallest, 268 mm. total length, collected June 29, 1961, 
has a yolk-sac scar and is slate gray above with a bright-yellow belly 
and the characteristic red subcaudal surface. Another, 377 mm., is 
similar to the juvenile ventrally but has a greenish-gray dorsum. The 
largest specimen, 478 mm. with a stub tail, has an orange-yellow 
belly that shades into coral red in the region of the vent and is also 
greenish gray dorsally. Its individual dorsal scales are mottled with 
black and have a black spot at the anterior end (pi. 4). In each of 
these three ringnecks, the belly is moderately to heavily spotted with 
black as in all other southwestern and Mexican specimens. 

LocaHty records: 

MCKixLEY CO.: 6 mi. S. Thoreau (UMMZ 120276, 122627, 122947); Valencia 
CO.: 6 mi. E. Grants (UNMCV 158). 

Comparative material examined : 

D. p. regalis. — New Mexico: bernalillo co.: SW>^, Sec. 22, TON, R4E 
(UNMCV 412); Jaral Canyon, Sandia Mts. (UNMCV 36); catron co.: 3 mi. 
N. Glenwood (BYU 13976); doxa ana co.: Mesilla Valley (USNM 22376); 
GRANT CO.: Mogollon Creek, above falls (UNMCV 159); Lincoln co.: 12 mi. 
NW. Carrizozo (UMMZ 66774); sandoval co.; E. Sandia Pueblo (UNMCV 
413); Pena Blanca (UNMCV 131); 25 mi. N. Albuquerque (UNMCV 163); 
SIERRA CO.: Lake Valley (ANSP 10786); torrence co.: Mountainair (UNMCV 
411); Arizona: cochise co.: 0.5 mi. SW. Portal (UMMZ 119029); Bisbee 
(CU 4694-5); Pinery Creek, Chiricahua Mts. (AMNH 80807); 2 mi. W. Ft. 
Huachuca (USMN 102201); Ft. Huachuca (USNM 21061); gila co.: Roosevelt 
(UNMCV 2698-9); final co.: Camp Grant (USNM 8019); navajo co.: Ft. 
Apache (USNM 8427); pima co.: Sabino Canyon, Santa Catalina Mts. (USNM 
62569); Sycamore Canyon, Baboquiviri Mts. (ANSP 17998); Otero Canyon, 
Baboquiviri Mts. (ANSP 17953); Ft. Buchanan (ANSP 3465); santa cruz co.: 
Madera Canyon, Santa Rita Mts. (ANMH 64428); 10 mi. N. Nogales (AMNH 
67253); yavapai co.: White River Canyon (USNM 10199); Camp Verde 
(AMNH 4202, 62240); Utah: san pete co.: Freedom (UMMZ 63999); utah 
CO.: S. Fork West Canyon (BYU 13775, 14168-9, 14672-3); Washington co.: 
Pine Valley (CU 3836); Mexico: chihuahua: Maguarichic (UMMZ 118929); 
6 mi. E. Barraganes (AMNH 73749); Norogachi (AMNH 73751); 10 mi. W. 
San Francisco del Oro (BYU 14251); 7 mi. SW. Pacheco (MVZ 46685). D. p. 
dugesi. — Mexico: hidalgo: 8 mi. N. Pachuca (UIMNH 17635); jalisco: 
10 mi. W. Guadalajara (UIMNH 46578, 47866) ; Guadalajara (AMNH 3711- 
12); W. Ocotlan (AMNH 19723); mexico: (AMNH 62827); 3 mi. S. Villa 
Obregon (UMMZ 99540); michoacan: 2 mi. E. Morelia (AMNH 62827); 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 307 

15 km. W. Morelia (UIMNH 17634); 5 mi. E. Lake Patzcuaro (UIMNH 17636); 
nayarit: La Labor (AMNH 19724); 7 mi. S. Compostela (AMNH 80590). 
D. p. regalis x diigesi. — Mexico: chihuahua: 2 mi. E. Cerocahui (BYU 14243); 
DURANGo: 20 mi. NW. Los Coyotes (UAIMZ 102527); 40 mi. SW. Cd. Durango 
(ISM uncataloged). D. p. regalis x arnyi. — New Mexico: eddv co.: Walnut 
Canyon, Guadalupe Mts. (CCNP 2197, 2201 ; UMMZ 122949) ; Texas: brewster 
CO.: near Alpine (CU 869, 883); near Panther Pass, Chisos Mts. (UNMCV 
2701); Upper Green Gulch, Chisos Mts. (UNMCV 2700); culberson co.: 
McKittrick Canyon, Guadalupe Mts. (UMMZ 121825, 122948, 122955; BCB 
8702). 

Masticophis flagellum Uneatulus Smith 

Since the description of M. f. Uneatulus (Smith, 1941), in which the 
specimen from Lagmia, Valencia Comity, was indicated as a paratype, 
only one additional individual, to my knowledge, has been collected 
in the Zuni region. It comes from the Plains Life Belt at 6300 feet, 
where narrow-leaved yucca and one-seed junipers are prevalent. This 
coachwhip snake may reach the approximate northwestern limit of 
its New Mexico range at the eastern end of the Zunis. It is presently 
unknown in McKjnley and San Juan Counties, although Maslin 
(1959, p. 57) saw what he presumed to be AI. flagellum in adjoining 
Montezuma Count}", Colorado. 

The recent specimen, an adult male, 1070 mm. in total length, 
has longitudinal dark lines on the dorsal scales but retains dark 
anterior cross bands and lacks the diagnostic red subcaudal color. 
Its subspecific allocation is, therefore, tentative. I must agree with 
Wright and Wright (1957, pp. 437-438), who remarked on the difficult 
southwestern members of the M. flagellum complex and considered 
Smith's contribution (1941, pp. 394-397) as tentative. 

Locahty records : 

VALENCIA CO.: Laguna (USNM 4388); 8 mi. E. Grants (TNHC 4487). 

Masticophis taeniatus taen ia tus Hallowell 

Smith and Taylor (1950b, p. 359) apparently overlooked historical 
circumstances in restricting the type locality of this whipsnake to 
Shiprock, San Juan County, New Mexico. The actual type locahty 
cannot be determined exactly, but it can be clarified by the following 
facts. S. W. Woodhouse, accompanying the 1851 Sitgreaves Expe- 
dition, collected the form described by Hallowell (1852, p. 181) as 
Leptophis taenita [sic] and later stated by him (Hallowell, 1854, p. 134) 
to have come from "New Mexico, west of the Rio Grande." Wood- 
house came no closer than 100 miles of Shiprock in his journey from 
Albuquerque to the Zuni Pueblo, but he could have taken M. t. 
taeniatus anywhere in the Rio Grande basin between El Paso and 
Zuni (Hallowell, 1854, p. 147). Apparently he did not collect snakes 
west of Zuni, for he was incapacitated partially by a bite from Crotalus 
viridis in that locality. 

768-287—65 5 



308 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ns 

In color and scutellation the few Zuni specimens do not deviate 
appreciably from M. t. taeniatus as defined by Ortenbm-ger (1928, 
pp. 25-35) ; however, two juveniles, 525 and 672 mm. in total length, 
lack the usual reddish suffusion on the subcaudal surface and have 
clear, undivided, dorsolateral white stripes on the posterior half 
of the body. Three individuals were collected or observed in the 
piny on- juniper savanna, and one came from the cholla-juniper asso- 
ciation. Judging from sight records, this whipsnake is more abundant 
in open Roughlands envkonments than the number of museum 
specimens indicates. The species is particularly adept at escaping 
capture. 

Locality records: 

VALENCIA CO.: 5 mi. E. Grants (CU 3052); 10.4 mi. S. Grants (CU 5602); 
MCKINLEY CO.: Nutda (USNM 8432). 

Salvadora grahamiae grahamiae Baird and Girard 

Records of S. g. grahamiae on the southeastern edge of the Zuni 
Mountains and at Santa Rosa, Guadalupe County, New Mexico 
(Bogert, 1939, p. 189), provide the northernmost stations for this 
species in the United States. Mountain patch-nosed snakes were 
reported previously from Valencia County by Charles M. Bogert 
(see Wright and Wright, 1957, p. 649), who informed me (in litt.) 
that he examined a specimen taken one mile east of Laguna. 

The present Zuni specimens are from 6400 feet in a choll a- juniper- 
dominated section of the Plains-Roughlands continuum. Both have 
a high number of ventral scales, 199 and 196 for the male and female 
respectively; subcaudals are 100 and 97, dorsal scale rows are 19-17-13, 
and the labials are 9/9 except for 8 in the left upper series of the 
female. The smaller male has no trace of lateral dark stripes, and 
its dorsal stripes disappear on the basal third of the taU. The female 
has definite, dark lateral stripes that become diffuse at the anus. 

Hartweg (1940) noted that one of two specimens from near Mim- 
bres, Grant County, New Mexico, lacked the lateral stripes, as did 
specimens from Trans-Pecos, Texas, and southeastern Arizona. 
Stebbins (1954, p. 501) stated that Bogert had seen intermediates 
between S. g. grahamiae and S. g. lineata (sometimes considered 
a distinct species) in southern Chihuahua and Durango, Mexico. 
I examined a specimen (UMMZ 118446) from near Sombrerete, 
Zacatecas, Mexico, that was also intermediate. Thus, it seems 
that the two races intergrade over an unusually broad area, or that 
polychromatism exists; I favor the latter explanation. 

Locality records: 

VALENCIA CO.: 10.4 mi. S. Grants (CU 5655). 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 309 

Pituophis melanoleucus affinis Hallowell x P. m. deserticola Stejneger 

In the Zuni region, gopher snakes present a situation resembhng 
the one already described for Phrynosoma douglassi. Although the 
type locality of P. m. affinis is probably about five miles from the 
Pueblo of Zuni (see Woodhouse, 1854, p. 34), the gopher snake popu- 
lation inhabiting this area actually contains intermediates between 
P. m. affinis and P. m. deserticola. Ivlauber (1947, p. 41) reviewed 
the circumstances of Hallowell's (1854, p. 146) original restriction of 
type locaHty to "near the Zuni River, New Mexico." Smith and 
Taylor (1950b, p. 359) further restricted it to Zuni, McKinley County, 
New Mexico, apparently overlooking Woodhouse's statement con- 
cerning the provenance of the type of P. m. aijinis. 

When Klauber (1947, p. 44) noted that P. m. affinis from the 
Flagstaff-Gallup area was atypical, he seemingly did not consider 
that intergradation between it and P. m. deserticola might extend that 
far south. Instead, he believed the blend zone to be in the vicinity 
of the Hopi villages east to Canyon de Chelly, Arizona, on the basis 
of intermediate specimens available to him. It now appears that the 
P. m. affinis x P. m. deserticola zone is much broader. Considering the 
characters cited by Klauber (1947), examples of both P. m. affinis and 
P. m. deserticola have been taken in San Juan and McKinley Counties 
(Gehlbach, 1956, p. 370), and intermediates come from the Zunis and 
33 miles southeast of Laguna, Valencia County (UMMZ 121490). 
Thus, at least part of the intergrade area extends south and east of 
the San Juan Basin. 

Zuni specimens are intergrades on the basis of color and body and 
tail blotch counts (table 9) . These characters are intermediate when 
compared to those given by lOauber (1947, table 4) for P. m. affinis 
and P. m. deserticola. With the exception of two individuals, reddish- 
brown anterior dorsal blotches predominate in the sample, but the 
influence of P. m. deserticola is invariably present in the black tail 
blotches. Two specimens, one with reddish and one with black 
anterior blotches, have these markings interconnected so that the 
light interspaces appear as isolated spots. Specimens least typical 
of P. m. deserticola, in having the highest number of dorsal blotches, 
have black blotches, thus indicating their intermediate status. In 
view of the sample as a whole, intergradation in every character may 
be observed, but P. m. affinis influence is probably the strongest 
overall. 

Only Thamnophis elegans and Crotalus viridis were more evident 
than gopher snakes in the Zunis. P. melanoleucus was found usually 
in the lower two life belts, being least abundant in Roughlands 
associations. More specimens were taken and observed in saltbush- 
sage than in any other vegetational type. In the Roughlands life 



310 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

belt, gopher snakes were encountered in open pinyon-juniper savannas. 
Two individuals came from a choUa-juniper section of the Plains- 
Roughlands ecotone, where they were secreted in pack rat nests 
built among Indian Ruins. A juvenile, 385 mm. total length, col- 
lected July 23, 1957, at 7100 feet, was undoubtedly a hatchling as it 
had a fresh yolk-sac scar. 

Of the variety of food items palpated from newly caught specimens, 
Sceloporus undulatus, Peromyscus species, Neotoma species, Thomomys 
bottae, and various passerine birds predominated. One gopher snake, 
extracted by the tail from a gopher burrow at 10:35 a.m., was 
coiled tightly around an adult Thomomys bottae. Juveniles sometimes 
fed upon lizards in the manner of a garter snake, i.e., without first 
constricting their prey. 

Locality records: 

MCKINLEY CO.: 4 mi. S. Thoreau (CU 5611); 4 mi. NNE. Prewitt (CU 5589); 
Ft. Wingate (USNM 8658, 16766-7); Black Rock (CWGF); Valencia co.: 6 mi. 
SE. Grants (CU 5612); 8 mi. SE. Grants (UMMZ 8G632); 16 mi. S. Grants 
(UMMZ 86631); 5 mi. SSW. U. S. Rt. 66 on N. M. Rt. 117 (CU 5769); 2.5 mi. 
NE. Ramah (CU 5454); 2 mi. W. El Morro Nat. Mon. (CWGF); El Morro Nat. 
Mon. (CWGF). 

Hypsiglena torquata texana Stejneger x //. t. ochrorhyncha Cope 
X H. t. loreala Tanner 

In New Mexico, as throughout the Southwest, some subspecies of 
H. torquata seem to me to be poorly defined. This fact and the paucity 
of specimens from the Zuni region demand that the present intergrade 
zone be considered provisional. Stebbins (1954, p. 492) has mapped 
this zone, utilizing subspecies boundaries defined by Tanner (1944). 
I Hkewise follow Tanner (1944) but with reservation, in light of the 
preceding discussion. Bogert and Oliver (1945, pp. 378-381) and 
others have presented evidence for lumping the species H. ochrorhyn- 
cha under H. torquata. 

The Zuni snake, a male, has 172 ventrals, 47 subcaudals, 21-21-17 
dorsal scale rows, 1-1 loreals, and approximately 49 large, dark body 
blotches, all characteristic of H. torquata texana, while the Los Lunas 
specimen, with about 52 body blotches, resembles H. torquata texana 
in color-pattern and H. t. loreala or H. t. ochrorhyncha in having 15 
dorsals posteriorly. UNMCV 79 is also intermecfiate; it has 2-2 loreals 
as in H. t. loreala, 16 posterior dorsals, and about 69 small dorsal 
blotches, similar in size (involving less than 20 scales) to those of 
H. t. loreala or H. t. ochrorhyncha. Ventral scale counts of the three 
specimens, 165, 172, and 177, approximate those of H. torquata texana 
or possibly H. t. ochrorhyncha rather than H. i. loreala (see Tanner, 
1944, table 2)s 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 311 

Features in common include the light-tan ground color and medium- 
olive or brown blotches ringed in interrupted fashion by dark-brown 
spots. Some dorsal blotches are broken into a zigzag row of separate 
or annectant spots as in other spotted night snakes examined from 
New Mexico. The Hght-tan coloration differs, however, from the 
medium gray manifest in H. torquata from limestone areas in southern 
New Mexico. Both UNMCV 79 and CU 5067 have continuous neck 
bands, a feature found in three of seven specimens at hand from 
southern New Mexico. A complete band is presumably less frequent 
in H. torquata texana than inH. t. loreala or H. t. ochrorhyncha (Tanner, 
1944). 

The single Zuni specimen was taken beneath a tan sandstone slab 
(Wingate) near the lower edge of the pinyon-juniper association at 
6800 feet. Large boulders, steep-sided canyons, and such reptiles as 
Urosaurus ornatus and Crotaphytus collaris typified the terrain. 
Another individual was seen in a nearly identical situation at 7200 
feet, two miles south of Thoreau, McKinley County, but escaped into 
a deep crevice. 

Locality records: 

MCKINLEY CO.: 3 ini. NE. Thoreau (CU 5067). 

Comparative material examined: 

H. torquata subspecies. — New Mexico: Valencia co.: Los Lunas (USNM 
107347) ; san juan co.: Chaco Canyon Nat. Mon. (UNMCV 79) ; eddy co.: Carls- 
bad Caverns Nat. Park (UMMZ 86163, 121785; CCNP 2409, 2416); catron co. : 
1 mi. N. Glenwood (UNMZ 78231); 13 mi. SE. Glenwood (UMMZ) 78232-33). 

Crotalus atrox Baird and Girard 

A characteristic member of southwestern desert and desert-grassland 
herpetofaunas, C. atrox is another species that probably reaches the 
northwestern limit of its New Mexico range in the Zuni region. Here 
it is found only in open plains situations but may range into the 
Plains-Roughlands continuum. Ecological restriction to the cholla- 
juniper association was observed south of Grants, where diamondback 
rattlers were found between the lava flow and edges of surrounding 
uplands. This snake was absent from the high soil-covered malpais 
near El Morro National Monument, where C. viridis was abundant; 
a similar situation was observed on the rough Grants malpais. A 
large adult, collected by William L. Chenoweth on October 17, 1957, 
was dead on the road in the short-grass association near Prewitt, when 
the air temperature was about 55° F. 

Yarrow (1875, p. 529) first recorded C. adamanteus atrox from Ft. 
Wmgate; Cope (1900, p. 1163) and VanDenburgh (1924, p. 227) 
mentioned this locality, the latter placing it in McKinley County. 
While diamondback rattlers occur in southeastern McKinley County, 



312 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

it is doubtful that the specimen in question, if identified correctly, 
came from the present Ft. Wingate, which is located in the pinyon- 
juniper association. On the basis of known distribution, it probably 
originated near San Rafael, Valencia County, where Ft. Wmgate 
formerly was located (Hoffmeister, 1951, p. 34); however, it is possible 
that the specimen was misidentified, for it was sent alive to the 
U.S. National Museum by R. W. wShufeldt and, seemingly, was 
never preserved (D. M. Cochran, in Utt.). 
Locality records : 

MCKINLEY CO.: Prewitt (UMMZ 79308); 8 mi. ESE. Prewitt (CU 5884); 
VALENCIA CO.: 8 mi. SE. Grants (UMMZ 86633); 10.4 mi. S. Grants (CU 5591); 
11.5 mi. SSE. Grants (CWGF); San Rafael (CWGF), 

Crotaliis molossus tnolossus Baird and Girard 

Chenoweth (1950, p. 534) first indicated that the black-tailed 
rattlesnake might be present in the Zimi region. Previously, the 
northernmost New Mexico record wasLaguna (LMK 3197), although 
a smashed individual was seen on the road near Cubero, Valencia 
County, by Charles M. Bogert (L. M. Klauber, in litt.). This 
species may range somewhat further north. James S. Findley (pers. 
comm.) informed me that C. molossus has been reported by other 
observers in the Sandia Mountains, Sandoval County, and near 
Cuba, Rio Arriba County. 

The single specimen, found during the present survey, was dead 
on the road in Lobo Canyon at approximately 7000 feet on Mt. 
Taylor. Its anterior color pattern is typical of C. molossus from 
south-central New Mexico, but there are only 12 distinct dorsal 
blotches, the entire posterior half of the body being uniform olive 
brown. The specimen has 27 dorsal scale rows at midbody; other 
counts cannot be ascertained because of its damaged condition. 

Locality records : 

VALENCIA CO.: 8 mi. NE. Grants (CU 5868); Laguna (LMK 3197*). 
Crotalus viridis viridis Rafinesque x C. v. nuntius Klauber 

Prairie rattlesnakes in the Zuni region present an east-west inter- 
grading situation similar to that of Crotaphytus collaris. Specimens 
collected east of the Apache County, Arizona-McKinley County, 
New Mexico, border have been considered nominate C. viridis 
previously (Gehlbach, 1956, p. 371; Klauber, 1935, p. 85). In describ- 
ing C. V. nuntius of the Hopi Indian country, Klauber (1935) stated 
that easterly intergradation of it with nominate C. viridis was 
gradual over a broad zone. He considered specimens from Gallup 
closest to C. V. viridis in coloration but with low ventral scale counts. 

When compared with comprehensive data presented by Klauber 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 313 

(1956, table 2.7), a series of prairie rattlers from the Zunis approxi- 
mates C. V. nuntius in six of eight meristic characters: number of 
subcaudal scales and tail rings in males, nmnber of ventral scales 
and tail rings in females, and total scale row and body blotch counts 
(table 9). Of the remaining two, only the female subcaudal count 
favors assignment to C. v. mridis; the male ventral count is intermedi- 
ate. Further indications of affinity with C. v. nuntius are the number 
of midtail scale rows (11-15, x 12.3±0.21) and ground color, which 
is predominately tan or brown rather than greenish. 

Fifteen adults average 653 mm. in total length. The largest is 
926 mm. and is the only decidedly greenish individual collected or 
observed. A female, 620 mm. total length, gave birth to six young, 
measuring 212-228 (x 219) mm., on September 21, 1957. Klauber's 
(1956, table 4.1) data for average size at bhth indicate that these 
young are approximately intermediate between C. v. viridis and 
C. V. nuntius. 

It is not surprising to note the light-tan to yellow ground color of 
C. V. concolor in a few Zuni specimens. This feature has been reported 
in San Juan County, New Mexico (Gehlbach, 1956, p. 371). Inter- 
grades between C. v. viridis, C. v. nuntius, and C. v. concolor probably 
occur in the San Juan Basin, but this is difficult to demonstrate since 
C. V. concolor is distinguishable from C. v. nuntius only by means of 
coloration. 

Two prairie rattlers, UMMZ 121413 from Canyon de Chelly 
National Monument, Apache County, Arizona, and UMMZ 121412 
from White Canyon, San Juan County, Utah, are pertinent to this 
discussion. Both are small adults, 540 and 485 mm. in total length 
respectively, alilve in their pinkish-brown coloration. The Arizona 
specimen differs by having white borders around its anterior dorsal 
blotches and head markings and a dark-brown stripe across the snout. 
The Utah snake could be C. v. nuntius x C. v. concolor in view of its 
color, pattern, and locality, but several of the larger Zuni specimens 
also lack white-bordered markings. Legler (1960, p. 182) states that 
dorsal markings are obliterated gradually with age in C. v. concolor. 
Apparently C. v. viridis does not penetrate northwestern New Mexico 
and adjacent Arizona and Utah as mapped by Klauber (1956, fig. 2.6). 

Prairie rattlesnakes inhabit a ^vide range of Zuni environments. 
While many individuals were taken on lava flows and in the short-grass 
and saltbush-sage associations, a smaller number were collected in the 
Roughlands Life Belt. One was found in the ponderosa pine associa- 
tion of the lower Montane Life Belt. C. viridis seemingly was 
outnumbered by C. atrox in the lower, more open grassland, especially 
that dominated by cholla cactus. Juveniles were uncommon in the 
Plains Life Belt, suggesting that they might be born in the vicinity 



314 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

of hibernation sites located in rocky uplands. Both juveniles and 
adults were often active during early morning and late afternoon hours; 
they contained recently ingested lizards as well as rodents. Two 
taken at 9:00 and 10:35 a.m., July 7, 1956, at 7200 feet had eaten 
Sceloporus undulatus. 

Jon A. Peterson and field companions described (in litt.) a "combat 
dance" involving two large individuals at 9:00 a.m., July 14, 1959, 
near Tinaja, Valencia County. The rattlesnakes, each about three 
feet long, were situated in bright sunlight in the open grassland of 
the Miocene lava plain at 7400 feet. A few junipers and pinyons 
were nearby. The snakes were twisted about each other anteriorly 
in a position vertical to the ground. After several minutes of observa- 
tion (photographs were taken, which I have seen), the snakes were 
disturbed; they were not collected. 

Locality records : 

MCKINLEY CO.: 4 mi. S. Thoreau (CWGF) ; 6 mi. S. Thoreau (CU 5046, 5446, 
5603); 10 mi. N\V. Thoreau (CU 5673); 5 mi. NE. Thoreau (CU 5674); 4 mi. 
NNE. Prewitt (CU 5457, 5633); Prewitt (UMMZ 79392); Ft. Wingate (USNM 
8399); 2.5 mi. N. Upper Nutria (KUMNH 45765); 11 mi. W. Ramah (UMMZ 
122882); Valencia co.: 1.5 mi. S. Grants (UMMZ 86635); 8 mi. SE. Grants 
(UMMZ 86634) ; 10.4 mi. S. Grants (CU 5590) ; 25 mi. S. Grants (MCZ 62267*) ; 
Tinaja (CU 5604) ; Acoma Pueblo (USNM 44483) ; El Morro Nat. Mon. (CWGF). 

Species of Questionable Occurrence 

Certain amphibians and reptiles that may occur in the Zunis are 
currently unrecorded. This undoubtedly obtains for medium to small 
colubrid snakes in view of the late discovery of Diadophis pundatus 
in the region. Arizona elegans, Elaphe guttata, Lampropeltis doliata, 
and Opheodrys vernalis are present in north-central New Mexico and 
southwestern Colorado or adjacent Utah and eventually may be 
collected in or near the Zuni region. The latter two species can be 
presumed to occur on the basis of observations by local residents. 
A. elegans, collected near Farmington (A. Harris, in litt.), is the only 
one of these snakes recorded from San Juan County, New Mexico: 
it has been taken near Madrone, Valencia County (UNMCV 407). 

Sceloporus magister, Crotaphyty.s wislizeni, and Cnemidophorus 
tigris also occur in north-central New Mexico and southwestern 
Colorado. The latter two lizards and Heterodon nasicus are charac- 
teristic of a type of sandy habitat seemingly lacking in the Zunis. C. 
wislizeni is present near Laguna (USNM 4274), but C. tigris has been 
taken no closer than nine miles south of Shiprock (AMNH 77530) 
and Blanco, San Juan County, New Mexico (A. Harris, in litt.). 
H. nasicus was collected between Laguna and Correo, Valencia 
County (UNMCV 391). S. magister is known from Montezuma 



HERPETOLOGY OF ZUNI MOXJNTAINS — GEHLBACH 315 

County, Colorado (Masliii, 1959, p. 30), and Los Lunas, Valencia 
County (Smith, 1949, p. 156); its apparent absence in the Zunis 
is enigmatical. 

A nmuber of species present in Chihuahuan desert-grassland in 
extreme eastern Valencia County might be found as far west as Laguna 
but cannot be expected on the Colorado Plateau. Of these, Phryno- 
soma modestum (UNMCV 1170) and Cnemidophorus tessellatus 
(UNAICV 1169) have been taken near Correo. Semiaquatic forms 
such as Thamnophis sirtalis and Chrysemys picta seem more closely 
restricted to the Rio Grande. T. sirtalis was reported from Los 
Lunas, Valencia County, by Fitch and Maslin (1961, p. 299), and a 
relict colony of C. p>icta (UMMZ 64916-17) also exists there. That 
turtles were formerly present in the Zunis is suggested by paintings of 
them on Zuni Indian pottery (Whipple, 1856, p. 66).^ 

Hyla eximia {=H. wrightoruml) listed from Nutria (AIcKinley 
County?) New Mexico, by Yarrow (1875, p. 524) has not been col- 
lected during the present study. One of the two original specimens 
(USNM 8508; the other apparently is lost) was examined, but identity 
could not be verified owing to its poor state of preservation. Also, 
P. modestum and Bufo cognatus, recorded from Cottonwood Gulch by 
Chenoweth (1950), have not been rediscovered despite intensive field 
work in and around that locality. Since these records are based on 
one specimen each and the locality is the site of a boys' summer 
camp, the records are thought to represent introduced individuals. 

Zoogeography 

The Zuni region is situated on the southeastern edge of the Colorado 
Plateau adjacent to the northeastern limits of the Basin and Range 
physiographic province. Its zoogeographic miportance is demon- 
strated by the fact that eight amphibians and reptiles exhibit limited 
gene flow between intergrading subspecies that are associated better 
with the Colorado Plateau and other northern areas or the Basin and 
Range. It is similarly important that six species reach their northern 
limits of range in the area. To facilitate discussion of these regional 
patterns, the Zuni herpetofauna may be arranged according to present 
centers of distribution (table 11); nevertheless, interpretation of past 
events and causal factors in distribution rests entirely on paleoeco- 
logical inference in the absence of local fossil records. 



7 Some shells of Chrysemys picta (AMNH 19888, 20580) from the pueblo ruins 
of Hawikuh, 12 miles southwest of Zuni, Valencia County, recently came to my 
attention. They have not been dated with precision, but it is known that 
Hawikuh was abandoned around 1670 A. D. 



316 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 11. — Present centers of distribution of the Zuni herpetofauna 
Transcontinental North America 



Ambystoma tigrinum 
Bufo woodhousei 
Pseudacris triseriata 
Rana pipiens 



Sceloporus undulatus 
Diadophis punctatus 
Masticophis flagellum 
Pituophis melanoleucus 



Western North America 



Scaphiopus inter montanus 
Crotaphytus collaris 
Sceloporus graciosus 
Uta stansburiana 
Urosaurus ornatus 
Phrynosoma douglassi 
Cnemidophorus velox 



Mexican Plateau 



Scaphiopus hammondi 
Bufo punctatus 
Hyla arenicolor 
Holbrookia maculata 



Scaphiopus bombifrons 



Great Plains 



Eumeces multivirgatus 
Thamnophis elegans 
Masticophis taeniatus 
Hypsiglena torquata 
Crotalus atrox 
Crotalus viridis 



Thamnophis dorsalis 
Salvadora grahamiae 
Crotalus molossus 



Eumeces obsoletus 



Patterns of Distribution 

Three widespread species, Sceloporus undulatus, Urosaurus ornatus, 
and Hypsiglena torquata, contain one geographic race restricted to the 
greater Colorado Plateau region and another to the southeastern 
Basin and Range province. These intergrade in the Zunis and else- 
where along the southern border of the Plateau.^ Bujo woodhousei, 
Pseudacris triseriata, Uta stansburiana, Eumeces multivirgatus, and 
Pituophis melanoleucus, from the same distributional groups (trans- 
continental and w^estern North America), have similar patterns of 
restricted gene exchange, but their northern subspecies are m no sense 
limited to the Plateau. It is significant that a past or present break in 
the gene flow of these eight eurytopic forms coincides with the Colorado 
Plateau-Basin and Range border. 

Scaphiopus bombifrons and Eumeces obsoletus, the two Great Plains 
species, and Salvadora grahamiae and Crotalus molossus, centered on 
the Mexican Plateau, are at or near the northern limits of the south- 
western portions of their range in the Zunis. They have not pene- 
trated Colorado Plateau highlands except along a few major river 
valleys. The five other species allied with the Mexican Plateau range 
somewhat further north but not beyond the upper Colorado River 

8 Holbrookia maculata and Crotalus viridis also may fit this pattern in part, but 
the Colorado Plateau subspecies involved are defined inadequately at present. 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 317 

basin. Crotalus atrox and possibly Masticophis flagellum also reach 
northern limits in the Zuni region. They are the only two species 
from the transcontinental and western North America groups that 
inhabit semiarid lowlands south of and, in the case of M. flagellum, 
east and west of the Colorado Plateau. 

No species of reptile or amphibian present in the Zunis is restricted 
to the Colorado Plateau, and none are definitely limited to it at their 
southern edge of range. Cnemidophorus velox is distributed largely 
within Plateau boundaries but also ranges into the Basin and Range 
province. The main distribution of Sceloporus graciosus apparently 
coincides with the southern border of the Plateau in New Mexico 
although relict colonies exist much farther south. Scaphiopus inter- 
montanus may reach its southern limits in the Zuni region, but is 
known too poorly in New Mexico to be of zoogeographic value. 
It appears, therefore, that Colorado Plateau environments have been 
more effective in limiting northward dispersal than in preventing 
movement southward. 

Physiographic continuity of the southern Plateau (Datil and 
Navajo Sections) across the Zuni region is in sharp contrast to the 
north-south break between the Plateau and Basin and Range south 
and, to some extent, southeast of this area. This situation does not 
favor east-west geographic differentiation but allows gene flow 
between subspecies that previously might have been better isolated. 
The intergradation patterns, thus, of Crotaphytus collaris and Crotalus 
viridis might be expected to differ in degree from those of the afore- 
mentioned species. There is reason to suspect that relatively smooth 
character gradients are present in the east-west pattern and that 
sharper breaks are involved in the more common north-south pattern. 

The remaining eurytopic species, Ambystoma tigrinum, Rana 
pipiens, Phrynosoma douglassi, Thamnophis elegans, Diadophis 
pundatus, and Masticophis taeniatus, are distributed throughout the 
Zunis, the Colorado Plateau, and bordering Basin and Range province 
with little apparent regional differentiation. While geographic 
variation occurs elsewhere, the demonstrated local barriers to dispersal 
are relatively imimportant in each of these species. 

Paleoecology and Dispersal 

As indicated by species adaptation and rather close restriction to 
modern environments, development of the Zuni herpetofauna — indeed, 
that of the entire Southwest — followed closelj^ upon the formation of 
recent topography and semiarid vegetation in the late Cenozoic. 
Differentiation of the Colorado Plateau from a relatively flat plain 
began in the Miocene, and epeirogenic uplift has continued into the 
Pliocene and Pleistocene, raising the Plateau above basins to the 



318 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

south and west (Hunt, 1956, pp. 73-86), The ranges of certain 
widespread species undoubtedly were fragmented first by these 
upheavals and by the climatic changes that followed. 

Dry, warm, middle Pliocene climates produced grassland and 
subdesert environments of great extent (Axelrod, 1948). Present 
regional adaptations such as time of egg-laying may have originated 
then in response to increasing aridity. It is probable that ensuing 
dry conditions permitted northward dispersal of amphibians and 
reptUes adapted to vegetative environments of the expanding Madro- 
Tertiary Geoflora and that montane species associated with forests 
of the Arcto-Tertiary Geoflora simultaneously were limited. Axelrod 
(1950, 1958) and Darrow (1961) have reviewed the history of these 
major geo floras. 

Many neontologists, including myself, believe that present plant 
and animal distributions were slightly to highly modified by violent 
climatic fluctuations during the Pleistocene. Considerable modifica- 
tion of the extent of plant communities characterized this epoch. 
Evidence presented by Antevs (1954), Clisby and Sears (1956), 
Leopold (1951a), Murray (1957), and Wendorf (1961) in New Mexico 
suggests that grassland and desert species withdrew southward at 
or about the time of glacial advance and that woodland forms then 
dispersed across former semiarid zones. 

Periglacial deposits are extensive on Mt. Taylor (Hunt, 1956, p. 
38) , indicating that the Zuni region was not excluded from Pleistocene 
climatic change. Glacial deposits have been recognized along the 
north, east, and south sides of the Colorado Plateau; these extend 
down to 7000 or 8000 feet in some areas (Hunt, 1956, p. 35). Martin, 
Sabels, and Shutler (1961, p. 115) postulated that during a cool- 
moist interval the pinyon-juniper savanna was displaced downward 
1700 feet in the Grand Canyon, Arizona, or, in terms of life zones, 
was 2000-4000 feet below its present lower limits. If Zuni life belts 
were uniformly lower by 4000 feet at such a time, as Antevs (1954) has 
estimated for lowered life zones at Santa Fe, then the present Plains 
Life Belt with its characteristic species did not exist. 

Zuni life belts probably fluctuated in a manner similar to those 
diagramed by Martin (1961, fig. 2) ; thus, plains species like Scaphiopus 
bombifrons and Holbrookia maculata were absent when cool-moist 
conditions favored heavy forests or open woodland at low elevations 
but may have been present prior to such forestation. Coincidently, 
woodland-canyon species such as Hyla arenicolor or Eumeces multi- 
virgatus were provided new pathways for dispersal. The opposite 
situation existed during warm-dry intervals; hence, woodland forms 
have had the most limited gene flow in Hypsithermal time and 
presumably within the past century. While such "alternating 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 319 

genetic currents" are purely speculative, they are in accord with 
paleoecological and neoecological evidence. It is significant that 
the north-south patterns of Pleistocene clunatic change and local 
physiographic differentiation augment each other in affecting dispersal. 

If cool-moist intervals limited the northward movement of wide- 
spread, grassland species, they were equally potent in restricting 
desert and desert-grassland forms like Crotalus atrox. No great 
changes in drainage pattern have occurred on the Colorado Plateau 
since the late Pliocene (Hunt, 1956, p. 85), but the Rio Grande valley 
of southern New Mexico did not exist as a lowland pathway for 
north-south dispersal prior to early mid-Pleistocene time (Ruhe, 
1960). Are some species that reach their northern limits in the 
Zuni region, therefore, postglacial invaders? Dowhng (1956) accounted 
for the presence of similar xerically adapted species in Arkansas by 
postulating recent immigration from the Great Plains during a 
Holocene arid interval. 

What about the distribution of such vagile forms as Amby stoma 
tigrinum and Thamnophis elegarts? It seems possible that at least 
some of them remained locally at lower elevations on the Colorado 
Plateau during the Pleistocene cHmatic shifts. This is suggested by 
Tihen's description (1942) of neotenic A. tigrinum in Great Plains 
glacial deposits and typical metamorphosed individuals in interglacial 
sediments. Many of the other widespread species are equally 
adaptable and some hke Phrynosoma douglassi, in addition to A. 
tigrinum and T. elegans, are relatively abundant in both Montane 
and Plains Life Belts. 

Summary 

Between 1951 and 1961, the systematics, ecology, and life history 
of 30 species of amphibians and reptiles were studied in the Zuni 
Mountains region, McKinley and Valencia Counties, New Mexico. 
This area, characterized by extremes of temperature and midsummer 
rainfall, contains Plains, Roughlands, and Montane Life Belts, and 
extensive lava flows. Elevations range from 5794 feet at Laguna on 
the eastern end of the study area to 11,389 feet at the top of Mount 
Ta3^1or. Unfortunately, man has drastically altered natural environ- 
ments. Drought pervades the recent climatic history of the region. 

Of 19 common species, no two have the same vertical distribution 
or relative abundance. Five lizards of similar adult size have different 
morphological and physiological adaptations and occupy different 
habitats. In the Iguanidae, scansorial species such as Urosaurus 
ornatus have more lamellae than terrestrial species like Phrynosoma 
douglassi. Oviposition in Holbrookia maculata, Sceloporus undulatus, 
Sceloporus graciosus, and U. ornatus seems to be correlated with 



320 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

the onset of summer rainfall. Scaphiopus hammondi, Crotaphytus 
colJaris, and U. ornatus are darkly colored on lava. Similarly, 
H. macvlata and S. graciosus are very light on white sand, but none 
of these species exhibit geographically unique coloration. 

Scaphiopus bombifrons, Scaphiopus intermontanus, Eumeces obsoletus, 
Salvadora grahamiae, Masticophis flagellum, Crotalus atrox, and 
Crotalus molossus reach their hmit of range in the Zuni region. Sub- 
specific intergradation occurs in Bufo woodhousei, Pseudacris triseriata, 
Crotaphytus collaris, Sceloporus undulatus, Uta stansburiana, Urosaurus 
ornatus J Eumeces multivirgatus , Pituophis melanoleucus , Hypsiglena 
torquata, and Crotalus viridis. In certain cases, distributional patterns 
may have been modified by climatic changes and the Colorado 
Plateau-Basin and Range physiographic break located immediately 
south and southeast of the Zuni region. The Plains Life Belt 
apparently was eliminated during cool-wet periods of the Pleistocene, 
and woodland environments were restricted during warm-dry intervals. 

Ambystoma tigrinum stebbinsi and A. t. utahense are synonymized 
with A. t. nebulosum, Holbrookia inaculata ruthveni with H. m. ap- 
proximans, and Phrynosoma douglassi ornatissimum with P. d. hern- 
andesi. Diadophis regalis and D. dugesi are considered to be sub- 
species of D. punctatus; the ranges of these forms and the western 
limits of D. p. arnyi are clarified. Detailed accounts of variation in 
A. t. nebulosum, Cnemidophorus velox, and southwestern and Mexican 
Diadophis are presented. Environmental and ontogenetic variation 
are given special emphasis, and a graphic method for illustratmg the 
latter is offered in the discussion of E. multivirgatus. 

Addenda 

In preparing the account of recent environmental changes, I over- 
looked an important paper by Leopold (1951b) containing a highly 
pertinent summary of some nineteenth-century observations of Zuni 
environments. Also instructive are two photographs (Leopold, 1951b, 
fig. 1) of an area near Fort Wingate showing encroachment of sage- 
brush on grassland between 1901 and 1946. 

The revised manuscript was completed in December 1961. Since 
then, Smith and Wilhams (1962) have shown that the names Eumeces 
multivirgatus gaigeae and Thamnophis crytopsis are conserved by ap- 
plication of the nomen oblitum rule of the 1961 International Code 
of Zoological Nomenclature. 

McCoy (1962) corrected Maslin's record (1959) of Crotalus viridis 
viridis in Montezuma County, Colorado; he identified three specimens 
from Mesa Verde as C. v. nuntius. Additional support for intergrada- 
tion between the subspecies C. v. viridis, C. v. nuntius, and C. v. concolor 



HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 321 

in the San Juan Basin of northern San Juan County, New Mexico, 
and adjacent Colorado was offered by Dean and Stock (1961). 

New evidence for considering Diadophis dugesi as a subspecies of 
D. punctatus wsls presented by McCoy (1964), who recorded a female 
from Durango, Mexico. His specimen is intermediate between the 
subspecies D. p. regalis and D. p. dugesi in ventral count (212) but 
resembles the latter in number of subcaudals (68) and dorsal scale 
rows (17-17). As I have shown, number of ventral scales is the most 
diagnostic feature separating southwestern and Mexican races of 
ringneck snakes. 

Dean and Stock (1961) reported Crotaphytus collaris auriceps in 
extreme northern San Juan and Rio Arriba Counties, New Mexico 
(San Juan Basm). If C. c. auriceps is indeed recognizable, then Zuni 
specunens seem to show a slight relationship with this form based on 
labial counts. It would be desirable to pursue other meristic features 
that might distinguish C. c. auriceps. 

A brief report by Maslin (1962) on all-female species of Cnemidopho- 
rus includes C. velox. The possibility of parthenogenesis is suggested; 
and two males of C. velox, possibly the ones I examined and thought 
close to C. inornatus, are mentioned. Maslin also considers their 
allocation questionable. 

Harris (1963) contributed data on the ecological distribution 
of reptiles and amphibians in the San Juan Basin. He recorded 
Scaphiopus bombifrons and Cnemidophorus inornatus in San Juan 
County, New Mexico, and elaborated other records contained herein 
and attributed to him. That the taxonomic relationships of S. 
bombifrons with *S'. inteimontanis and C. velox with C. inornatus need 
investigation becomes increasingly apparent. 

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HERPETOLOGY OF ZUNI MOUNTAINS — GEHLBACH 327 

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328 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

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330 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

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332 PROCEEDINGS OF THE NATIONAL ]MUSEUM vol, iie 

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U.S. GOVtRNMENT PRINTING OFFICEil98B 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 116 1965 Number 3506 

REVIEW OF THE GENUS CERCERIS 

IN AMERICA NORTH OF MEXICO 

(HYMENOPTERA: SPHECIDAE) 



By Herman A. Scullen ' 



Introduction 



The wasp tribe Cercerini as recognized in this country (Muesebeck, 
Krombein, and Townes, 1951) consists of the two genera Cerceris Latreille 
(1802) and Eucerceris Cresson (1865). The former is well represented 
in aU continental areas while the latter is limited to North America. 

The earUest references to the genus Cerceris in North America were 
by Thomas Say when he published descriptions of the following species : 
jrontata Say (1823), deserta Say (1825) , fumipennis Say, (1837) and 
sexta Say (1837). Numerous species have been described since the 
publications of Say. In 1865 E. T. Cresson monographed the Family 
Philanthidea of North America, which at that time included the 
genus Cerceris and the new genus Eucercens. Cresson published over 
20 papers relating to Cercerini and described a total of 20 species 
recorded from north of the Mexican border. In 1912 Nathan Banks 
published his first paper on the genus Cerceris. This was followed by 
eleven additional papers. In all, Banks described 52 species of 



1 Professor Emeritus, Department of Entomology, Oregon State University, 
Corvallis, Oreg. Grants from General Research (Research Paper no. 469) under 
the Graduate School and from the National Science Foundation have assisted in 
these studies. 

333 



334 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Cerceris.noTth. of the Mexican border. In spite of the fact that many 
of Bank's species have proven to be synonyms, his contribution has 
been outstanding. The work of H. S. Smith and C. E. Mickel in 
Nebraska is worthy of special note. In describing new Mexican 
species, Saussm-e (1867) and Cameron (1890) included several species 
found subsequently to range north of the Mexican border. Numerous 
other workers have added one or more species to the present list. 

A summary of all contributors (with the number of species described 
and their principal publication dates in parentheses) is as follows: 
Say (4, 1823, 1825, 1837); Spinola (1, 1841); Dahlbom (2, 1845); 
Guerin (2, 1845) ; F. Smith (4, 1856) ; Cresson (20, 1865, 1872) ; Packard 
(1, 1866) ; Saussure (6, 1867) ; Taschenberg (1, 1875) ; Patton (1, 1880) ; 
Schletterer (1, 1887) ; Provancher (1, 1888) ; Cameron (4, 1890, 1904); 
Fox (2, 1893) ; Cockerell (1, 1897) ; Viereck (1, 1902) ; Viereck and Cock- 
erell (17, 1904); H. S. Smith (2, 1903); Rohwer (6, 1908, 1920); 
Banks (52, 1912, 1913, 1915, 1916, 1917, 1919, 1923, 1947); Mickel 
(15, 1916, 1917); Brknley (5, 1927, 1928, 1929). 

Fourteen new species and 12 new subspecies are described by the 
author in the present paper. 

Biology. — Members of the genus Cerceris are solitary in their 
nesting habits although several females usually are found nesting close 
together. The grouping of several nests in the same area has been 
considered to be the result of the suitability of soil conditions and the 
availability of food for both larvae and adults. Krombein's observa- 
tions (1936) on C. nigrescens F. Smith, however, indicate there are 
exceptions. Apparently the adults feed solely on nectar and pollen, 
while the larvae are fed beetles.^ Only a limited number of species of 
Cerceris have been studied carefully to determine their nesting and 
feeding habits. Most species of Cerceris use some species of weevil to 
feed their young. On the other hand, the females of Group II are 
known to use Buprestidae. Both species forming Group IV {macro- 
sticta Viereck and CockereU and jemurruhrum Viereck and CockereU) 
are known to use Tenebrionidae. Recently, Werner (1960) reported 
C. truncata Cameron using three species of Bruchidae in Arizona. C. 
convergens Viereck and Cockerell also has been recorded collecting 
Bruchus sp. in southern California. Several species of Cerceris have 
been found to collect ChrysomeUdae. 

One observation recorded by J. C. Bridwell in unpublished notes is of 
special interest. Females of C. fumipennis Say under observation at 
Clifton, Virginia, were found "to be storing their nests with adult 



2 Some European species of Cerceris are known to store solitary bees as food for 
their young. For a brief discussion and references, the reader is referred to Linslcy 
and MacSwain (1956, p. 72). The records of Cerceris collecting insects other than 
beetles in this country are open to question. 



WASP GENTJS CERCERIS — SCULLEN 335 

beetles of the genus Chlamisus, a Chrysomelidae resembling Bupresti- 
dae only in their hard integument and their metallic luster." This 
change of prey took place after Buprestidae ceased to fly. 

Two unpublished records (Princeton, N.J., July 5, 1948, by K. W. 
Cooper, and Winchester, Va., July 23, 1961, by Frank Kurczewski) 
of C. clypeata Dahlbom taking Lema trilineata Oliver, a Chrysomelidae, 
have come to the writer's attention. C. clypeata normally takes 
weevils. 

Noteworthy published contributions in the field of Cerceris biology 
north of INlexico are summarized briefly below. 

The Peckhams (1898, pp. 108-117) report observations made on 
C. clypeata Dalhbom, C. deserta Say, and C. nigrescens F. Smith. C. 
clypeata Dahlbom was seen taking Balaninus nasicus Say. C. 
deserta Say was reported taking Conotrcbchelus posticatus Boheman. 
From Peckham's statement that C. deserta Say "closely resembles 
clypeata," it would appear the observations were made on a species 
other than the one accepted as deserta Say by Cresson, the present 
writer, and others (see p. 478). The above beetles are Curculionidae. 
The prey of C. nigrescens F. Smith was not recorded. In 1900 (p. 90) 
the Peckhams published a brief note on C. fumipennis Say found 
nesting in Wisconsin. This wasp was found collecting Chrysobothris 
4-impressa Castelnaw. 

Hartman (1905, p. 66) in reporting briefly on the nesting habits of 
C. fumipennis Say indicates it was collecting the weevil Conotrachelus 
neocrataege} This species of wasp usually takes Buprestidae. 

Grossbeck (1912a, p. 135) reported briefly on observations made at 
Yaphank, Long Island., N.Y., in 1911 on nesting Cerceris fumipennis 
Say. He reported the wasp storing the following Buprestidae: 
Dicerca punctulata (Schonherr), Buprestis lineata Fabricius, and 
Chrysobothris floricola Gory. 

Rau and Rau (1918), reporting on field studies near St. Louis, Mo., 
record the activities of C. fumipennis Say (pp. 126-127) and C. 
finitima Ci-esson (p. 133). The former wasp was found using Argilus 
abjectus Horn (Buprestidae). The prey of the latter wasp was not 
recorded. It was found to have made a nest in the pith-chamber of a 
weed-stalk although the Raus had noted it earher entering holes in the 
ground as is usual for the genus. Rau (1928, pp. 325-341) reports 
detailed studies of C. raui Rohwer {=C.frontata raui Rohwer), which 
he found nesting at Jerseydale about thirty miles south of St. Louis, 
Mo. This wasp was found using Thecesternus humeralis Say and 
Lixus concamis Say. Rau here reports also on his studies of C. 
bicornuta Guerin, which was using three species of billbugs, Spheno- 

^ C. neocrataege is not a valid name in the genus Conotrachelus. Correct identi- 
fication of the material Hartman had is not possible. 



336 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

jahorus placidus (Say), S. zeae Walsh, and S. parwlus Gyllenhal. 
The cuckoo-wasp Hedychrum violaceum Brulle was seen entering the 
bicornuta burrow. On page 338, Rau reports that C. jinitima Cres- 
son "make nests in twdgs," but the present writer finds no other 
record of this or any other species of Cerceris nesting in twigs. 

In 1928 (pp. 205-206) Abbott reported observations on the nesting 
and prey of Cerceris architis Mickel (— C halone Banks), which he 
found nesting at Elgin, 111. This wasp was using Curculio nasicus 
Say as food for its young. 

In 1929 (p. 35) Cartwright reported C. bicornuta Guerin provision- 
ing its nests with the maize billbug Calendra maidis Chittenden. 
Cartwright in 1931 (pp. 269-270) reported some observations at 
Jocassee, S.C., on C. Jumipennis Say, which he found collecting the 
beetle Buprestidae. Specimens of these beetles found about the 
wasp's nesting area numbered 136 and represented 20 species that he 
listed by name. At the same time, he reported seeing a specimen of 
C. mandibularis Patton bringing in a grasshopper nymph. This 
unusual prey record needs verification. 

Strandtmann reported briefly in 1945 (pp. 311-312) observations 
of three species of Cerceris in Ohio. C. serripes Fabricius (=C. 
bicornuta Guerin) was reported collecting a weevil, Calandra sp.; 
C. rujinoda Cresson, the weevil Tychius picirostris (Fabricius) ; C. jinitima 
Cresson, a black flea beetle, Cheatocnema pulicaria Melsheimer. 

Krombein has taken advantage of the opportunity to study the 
biology of several species of Cerceris over the past years. In 1936 
(pp. 93-99) he reported observations made of C. nigrescens F. 
Smith at Buftalo, N.Y. He found this vv^asp provisioning its nest with 
Hyper odes delumbis GyUenhal and Sitona hispidula Fabricius. A 
sarcophagid (Senotainia trUineata Van der Wulp) was seen shadowing 
the wasp. In 1938 (pp. 1-3) Krombein reported subsequent observa- 
tions at Buffalo on C. nigrescens F. Smith, where the wasp was found 
to be using the beetles Sitona hispidula (Fabricius), Oymnetron 
antirrhini Paykull, Oymnetron sp., and Hyperodes delumbis (GyUenhal), 
aU of which are Curcuhonidae. 

In 1950 (pp. 147-149) Krombein reported studies of a colony of 
C. robertsonii Fox at Kill Devil Hills, N.C. This species of wasp was 
found provisioning its ceUs with a chrysomelid (Rhabdopterus picipes 
(Oliver)). Aletopia leucocephala (Rossi), a parasitic sarcophagid fly 
was reared from the wasp cell. Subsequent studies by the same 
worker in 1952 at Kill Devil Hills, N.C. (1953, pp. 113-121), showed 
this wasp to be collecting Cryptocephalus notatus Fabricius and Pachy- 
brachis dilatatus Suffrian (Chrysomehdae). At the same location he 
reported C. bicornuta bicornuta Guerin taking the following weevils: 
Calendra venata venata (Chittenden) and C. cariosus (OHver). A more 



WASP GENUS CERCERIS — SCULLEN 337 

extensive report ou the nesting habits of C. robertsonii Fox is given by 
Krombein in 1952c (pp. 288-294). This same worker (1956, p. 43) 
pubHshed a brief note on the nesting of C. atramontensis Banks at 
Dunn Loring, Fairfax Co., Va., in which he reported the wasp collect- 
ing the weevils Conotrachelus posticatus Boheman and C. naso LaConte. 

In 1958 (p. 110) Krombein reported observations on the nesting of 
C . jumipennis Say at Kill Devil Hills, N.C., where he found the wasp 
utilizing the buprestids Chrysobothris femorata (OUver) and Dicerca 
lurida (Fabricius) as food for its young. 

In 1958 Krombein found a colony of Cerceris jiavofasciata H. S. 
Smith nesting at Kill Devil Hills, N.C., in a vertical sand bank (1959, 
pp. 197-198). These wasps were found to be using as prey for their 
young the following chr3^someHd beetles: Cryptocephalus guttulatus 
OHver, C. mutabilis Melsheimer, Cryptocephalus sp. (probably quadri- 
maculatus Say), Bassareus clathratus (Melsheimer), Bassareus sp. 
(probably sellatus Suffrian), and Chlamisus sp. (probably plicata 
(LeConte)). A mutiUid, Dasymutilla nigripes (Fabricius), was reared 
from a cocoon of the wasp. 

In 1959 Krombein (1960a, pp. 75-77; 1960b, pp. 299-300) studied 
the nesting habits of C. bicornuta jidelis Viereck and CockereU and 
C. frontata frontata Say at Portal, Ariz. The former wasp was found 
to be storing a weevil, Eupagoderes sp.; the latter wasp was storing 
Cleonus pidvereus (LeConte) and Eupagoderes sp. 

In 1956 Linsley and MacSwain pubhshed extensive studies of the 
nesting and prey collecting habits of Cerceris californica Cresson, 
which they found at San Dimas Experimental Forest and Range Ex- 
periment Station, San Gabriel Mountains, CaUf. These workers 
found C. californica Cresson using a total of 24 species of Buprestidae 
as prey to feed their young. Considerable information also is included 
on the parasites of the wasp. These include Dasymutilla coccineohirta 
(Blake) and the following sarcophagids : Metopia leucocephala (Rossi), 
Amobia floridensis (To^vnsend), and Senotainia trilineata (Van der 
Wulp). This is the most extensive report on the biology of a single 
species of Cerceris. 

Wasbauer (1957, p. 131) published a brief note on the biology of 
C. athene Banks (= C. femurrubrum athene Banks) at Mecca, 
Riverside Co., Calif. The prey taken by this wasp was a tenebrionid 
{Eurymetopon rujipes Eschscholtz) . It is significant that C. macro- 
sticta Viereck and Cockerel!, also belonging to Group IV, is known to 
coUect Tenebrionidae in Colorado and Arizona. From Wasbauer's 
observations it was found that C. athene Banks is the female of C. 
femurrubrum Viereck and CockereU (see p. 436). 

Werner (1960, pp. 43-44) published his observations on a nest of 
Cerceris truncata Cameron found nesting in his own yard. This wasp 



338 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

was found to be storing its cells with four species of native Bruchidae 
(Mylabridae) : Algarobius prosopis (LeConte), Mimosestes protractus 
(Horn), M. amicus (Horn), and Neltumius arizonensis (Schaeffer). 
Rau (1933, pp. 164-165) also reports a wasp (Cerceris near mexicana 
Saussure) as collecting Bruchus. 

Krombein (1963, pp. 72-79) published his observations on nests of Cercer- 
is blakei Cresson nesting in Florida. This was found to be storing its 
cells mostly with the weevil Derelomus basalis LeConte but was also 
taking the weevils Limnobaris confusa Bocheman, Anthonomus 
sexguUatus Dietz, and Hyperodes sp. Less commonly taken were the 
chrysomelid Graphops floridana Blake and the tenebrionid Blapstinus 
interruptus (Say). 

Cazier presently is conducting careful studies into the biology of 
several species of Cercerini at Portal, Ariz. It is probable that some 
of his studies will be reported before this publication appears. 

Byers (1962, pp. 317-321) reports finding a colony of Cerceris halone 
Banks nesting at Mountain Lake Biological Station on Salt Pond 
Mountain near Pembroke in Giles Co., Va., in the summer of 1961. 
This wasp was collecting Curculio nasicus (Say) as prey for its young. 
The cuckoo wasp, Hedychrum violaceum Brulle, was observed entering 
the wasp nest. At least three species of sarcophagid flies were re- 
ported seen in the nesting area. 

Several of the above investigators have made it a point to determine 
if the beetle prey is killed by the sting of the wasps or if it merely is 
paralyzed. From the published reports it would appear there is no 
consistency in this matter even with an individual wasp or within the 
same species. 

Numerous floral visiting records are included on the labels of 
specimens studied by the present writer. These and his own observa- 
tions show that most, if not all, species of Cerceris have their prefer- 
ences as to flowers visited for food by the adults. Open types of 
flowers with a liberal supply of pollen seems to be the usual preference. 
When the collector or student of Cerceris is looking for specimens in the 
field, it is important to keep in mind this floral preference and to 
remember that the difi^erent species of Cerceris may be visiting different 
species of flora in the same locality at the same time. A record of the 
flora on which they are collected is desirable. The flora from which 
they may be collecting beetle prey may be quite different from the 
flora visited for food by the adults. 

Morphology and taxonomic characters. — (Plate 1) . The writer 
has found most of the descriptions of older species very inadequate. 
For this reason and to make the descriptions uniform, all former 
species have been redescribed. Illustrations have been provided 
wherever it seemed desirable to clarify characters used. 



WASP GENTJS CERCERIS — SGULLEN 339 

Color shades and color patterns that have been used extensively 
by earlier describers are used by the writer to a more limited extent. 
He has found the color markings of some parts are far more stable 
than those on other parts within the same species. Futhermore, the 
overall tendency for color markings to vary is different for different 
species. One of the most extreme cases of color variation within 
a species is C. calijornica Cresson. Since, in some cases, the color 
changes are associated closely with geographical distribution, it seems 
desirable to recognize subspecies. 

In the females the form of the clypeal process and clypeal border 
continue to be the most important characters to use in separating 
species. For the males, the relative width of the hair lobes on the 
clypeus is important in separating groups but not individual species. 
Denticles on the mandibles, which were seldom mentioned by earlier 
workers, have been found of value in numerous cases. Other char- 
acters seldom or never mentioned by former workers but found useful 
by the writer are the form and sculpturing of the tegulae, the sculptur- 
ing of the enclosure, the mesosternal process, and the form of the 
pygidium. The form of the pydigium has been of special value in 
separating females. Extensive studies have been made of the male 
genitalia in an effort to find more dependable characters to help in 
their identification. Noticeable differences are found when comparing 
the males of one group with those of another, but, when species are 
otherwise closely related, no usable differences have been found. 

In general, the writer has found it far more diflBcult to separate 
males than females. Even in some cases wherein the males definitely 
have been recognized by their association with known females, no 
characters as yet have been found that will separate them. Here is 
an important field of study in this genus that should be pursued. 

Natural subdivisions. — Structures and biological behavior show 
certain natural divisions within the genus Cerceris. In the present 
publication the writer recognizes five more or less distinct groups. 
Further study may show that some or all of these groups are sufficiently 
distinct to warrant recognition as subgenera. For the present, it 
seems desirable simply to separate them as groups. Other natural 
subdivisions, not herein recognized, may appear with fiu"ther studies. 
Future structural and field biological studies may help in separating 
distinct groups. Ungrouped species are arranged alphabetically 
separate from the groups. The characters of each group are presented 
under each heading. 

Material studied. — Through the kind cooperation of many 
persons, the writer has been able to study well over ninety percent of 
the material from north of Mexico now found in the collections of 
this country and also many collections in Europe. Over 40,000 



340 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

specimens have been made available to him. Type material in the 
museimas of this country has been studied on several occasions, and, 
with the aid of a grant from the National Science Foundation, the 
writer was able to study types from this hemisphere deposited in 
European institutions. 

The following institutions have loaned material for these studies: 

Academy of Natural Sciences of Philadelphia, Philadelphia, Pa. 

American Museum of Natural History, New York, N.Y. 

Arizona State University, Tempe, Ariz. 

Boston Society of Natural History, Boston, Mass. 

Brigham Young University, Provo, Utah 

Brooklyn Museum, Brooklyn, N.Y. 

California Academy of Sciences, San Francisco, Calif. 

Canadian National Collection, Ottawa, Canada 

Carnegie Museum, Pittsburgh, Pa. 

Centre de Recherches Agronomiques de L'Etat, Gembloux, Belgium 

Clemson College, Clemson, S.C. 

Colorado State University, Fort Collins, Colo. 

Cornell University, Ithaca, N.Y. 

Deutsches Entomologisches Institut, Berlin, Germany 

Emory University, Atlanta, Ga. 

Hancock Foundation, University of Southern California, Los Angeles, Calif. 

Harvard University, Museum of Comparative Zoology, Cambridge, Mass. 

Illinois Natural History Survey, Urbana, 111. 

Instituto Technologico y de Studios Superiores, Monterrey, Mexico 

Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium 

Iowa State University, Ames, Iowa 

Kansas State University, Manhattan, Kans. 

Los Angeles County Museum, Los Angeles, Calif. 

Louisiana State University, Baton Rouge, La. 

McGill University, Redpath Museum, Montreal, Canada 

Michigan State University, East Lansing, Mich. 

Milwaukee Public Museum, Milwaukee, Wise. 

Montana State College, Bozeman, Mont. 

Museum Histoire Naturelle, Geneva, Switzerland 

Museum National d'Histoire Naturelle, Paris, France 

Natural History Museum, San Deigo, Calif. 

Naturhistorisches Museum, Zoologische Abteilung, Vienna, Austria 

New Mexico State University, Las Cruces, N. Mex. 

North Carolina State Department of Agriculture, Raleigh, N.C. 

North Carolina State College, Raleigh, N.C. 

North Dakota State University, Fargo, N. Dak. 

Ohio State University, Columbus, Ohio 

Oklahoma State University, Stillwater, Okla. 

Oregon State University, Corvallis, Oreg. 

Pennsylvania State Department of Agriculture, Harrisburg, Pa. 

Pomona College, Claremont, Calif. 

Purdue University, Lafayette, Ind. 

Rockefeller Foundation, Mexico City, D.F., Mexico 

San Jose State College, San Jose, Calif. 

South Dakota State College, Brookings, S. Dak. 



WASP GENUS CERCERIS — SCULLEN 341 

Stanford University, Palo Alto, Calif. 

State Plant Board of Florida, Gainesville, Fla. 

Sul Ross State College, Alpine, Tex. 

Texas Agricultural and Mechanical University, College Station, Tex. 

United States National Museum, Washington, D.C. 

University of Alberta, Edmonton, Alberta, Canada 

University of Arizona, Tucson, Ariz. 

University of Arkansas, Fayetteville, Ark. 

University of British Columbia, Vancouver, B.C., Canada 

University of California, Berkeley, Calif. 

University of California, Davis, Calif. 

University of California, Los Angeles, Calif. 

University of California, Riverside, Calif. 

University of Colorado, Boulder, Colo. 

University of Georgia, Athens, Ga. 

University of Idaho, Moscow, Idaho 

University of Kansas, Lawrence, Kans. 

University of Kentucky, Lexington, Ky. 

University of Massachusetts, Amherst, Mass, 

University of Miami, Coral Gables, Fla. 

University of Michigan, Ann Arbor, Mich. 

University of Minnesota, St. Paul, Minn. 

University of Nebraska, Lincoln, Nebr. 

University of Utah, Dugway Proving Grounds, Tooele, Utah 

University of Wisconsin, Madison, Wis. 

University of Wyoming, Laramie, Wyo. 

Utah State University, Logan, Utah 

Washington State University, Pullman, Wash. 

West Virginia University, Morgantown, W. Va. 

Materials from the personal collections of the following persons 
have been loaned for study: Rollin W. Baker, J. Bequaert, George E. 
Bohart, Kichard M. Bohart, Duis D. Bolinger, E. R. Buckwell, 
Walter Carter, Margaret L. Cook, Kenneth W. Cooper, Richard Dow, 
R. R. Dreisbach, Carl D. Duncan, George R. Ferguson, H. T. 
Fernald, Roland L. Fischer, J. E. Gillaspy, C. H. Hicks, Harry A. Hill, 
Gordon A. Hobbs, Paul D. Hurd, H, E. Jaques, Stanley G. Jewett, 
Wyatt W. Jones, Karl V. Krombein, Roy Latham, Howard L. Mc- 
Kenzie, C. D. Michener, H. E. Milliron, Reginald H. Painter, H. B. 
Parks, L. P. Rockwood, Vmcent D. Roth, C. W. Sabrosky, P. H. 
Timberlake, and M. Wasbauer. 

Acknowledgments.— Financial assistance which has made possible 
studies in the tribe Cercerini over the past thirty years has come from 
two sources. Grants of variable amounts have been received for many 
years from the General Research Council, Oregon State University, 
Corvallis, Greg. In more recent years three separate and substantial 
grants have been received from the National Science Foundation. 
These latter grants have made it possible to take two extensive 
collecting trips into Mexico (1956, 1957), to make an extended trip 



342 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii6 

through the East to study types at several institutions (1958), and to 
study types in European institutions (1959). Financial assistance 
from both sources has made it possible to secure secretarial aid, 
laboratory help, and illustrative assistance. 

Over the years many individuals have assisted in maldng these 
studies possible. While it would be impracticable to name all who 
have contributed, a few should be indicated. The most outstanding 
has been Dr. Karl V. Ej-ombein of the United States Department of 
Agriculture. Over many years he has been the writer's principal 
advisor; fiu-ther, he has contributed much original information through 
his many field studies; and finally, he has freely devoted much of his 
time to checking the present manuscript with special attention to 
the keys. Without his devoted help it is doubtful if this paper 
could have been completed. Among the technical assistants who have 
given noteworthy assistance, four deserve special mention: Marybeth 
Buddenhagen, Helen Dickason, and Margaret Johnson, who helped 
in checking records and references as well as many other details 
associated with such an undertaking, and Margaret Hsieh, who is 
responsible for most of the illustrations. The writer also is deeply 
indebted to Dr. Paul O. Ritcher, Head of the Entomology Department, 
Oregon State University, for the use of facilities to continue these 
studies after retu*ement. Sincere appreciation is here expressed to my 
wife, Johanne, for many hours of assistance in proof reading. 

Key to the Species of Cerceris 

FEMALES 

1. Stigma dark amber or black; mesosternal process usually present; species 

mostly small Group I 

Stigma light amber; mesosternal process absent; species mostly medium to 
large 2 

2. Clypeus without a distinct medial elevation or process on the surface of the 

medial lobe; a pair of small tuberculi (sometimes very indistinct) on the 
surface of the medial clypeal lobe just above the border; entire border 

of the clypeus without denticles Group II 

Clypeus with a distinct medial elevated process (sometimes little more than 
a ridge but not just a convex area) on the medial lobe; distinct extensions 
or denticle-like processes on the clypeal free border 3 

3. Clypeal process with a lamella or lamellae on the free border . . Group III 
Clypeal process without a lamella or lamellae on the free border .... 4 

4. Pygidium not narrowing to an acute or near-acute base (sides may converge 

slightly basally) 5 

Pygidium narrowing basally to an acute or very narrow end 36 

5. Elevation on the medial lobe of the clypeus is cone shaped or with the distal 

end very much narrowed and without a distinct overhang 6 

Elevation on the medial lobe of the clypeus with a definite overhang; the 
sides of the process may diverge, converge, or be subparallel 7 



WASP GElSrUS CERCERIS — SCULLEN 343 

6. Entire medial lobe of the clypeus elevated into a cone flanked by single but 

less prominent pointed processes on each lateral lobe .... Group IV 

Clypeal elevation or process not flanked by smaller processes on the lateral 

lobes Group V 

7. Lateral apices of the clypeal process considerably prolonged, giving the 

appearance of horns or the points of a new moon 8 

Lateral apices of the clypeal process, if any, not greatly prolonged ... 12 

8. Secondary lateral apices on the sides of the clypeal process . alceste Mickel 
No secondary lateral apices on the sides of the clypeal process 9 

9. Small species, approximately 10 mm. in length .... dentifrons Cresson 
Species of medium to large size, 15 mm. or more in length 10 

10. Distinctly black with creamy white markings .... stigmosalis Banks 
Black with yellow markings 11 

11. Apical mandibular denticle subequal to or smaller than the other denticles; 

distance between apical and medial denticles about one-half mm., medial 
and basal denticles confluent; clypeal process horns very long, curved, 
and acute; no yellow on the 6th tergum (Central States) . . . sexta Say 
Apical mandibular denticle noticeably larger than the other denticles; 
distance between all denticles subequal; 6th tergum yellow to the sides 
of the pygidium (west of the Rocky Mountains and into Mexico) . 

tepaneca Saussure 

12. Clypeal process width subequal to or greater than the length 13 

Clypeal process noticeably longer than the width 27 

13. Clypeal process very short, little more than a curved carina 14 

Clypeal process length subequal to its width 15 

14. Length 13 to 15 mm.; band on tergum 2 much broader than other abdominal 

bands but deeply emarginate halone Banks 

Length 10 to 11 mm.; bands on terga 2 to 5 subequal deserta Say 

15. Clypeal process relatively flat above 16 

Clypeal process convex above 18 

16. Free margin of the clypeal process showing very little emargination, lateral 

apices not acute, minute medial denticle on the margin (Eastern States 

from Kansas to the New England States) banks! ScuUen 

Free margin of the clypeal process with considerable emargination, lateral 
apices acute, margin without a medial denticle 17 

17. Legs amber with no yellow markings (Texas) texana ScuUen 

Hindfemora largely yellow with amber at the apical end only (Central Plains 

States from North Dakota to Texas) . . . occipitomaculata Packard 

18. Sides of clypeal process subparallel 19 

Sides of clypeal process converging 21 

19. Creamy yellow markings (Northern States from coast to coast) . 

nigrescens nigrescens F. Smith 
Yellow markings 20 

20. Somewhat more robust; lateral apices of clypeal process less acute (California 

and bordering states) nigrescens munda Mickel 

Lateral apices of the clypeal process very acute (Northeastern States) . 

nigrescens arelate Banks 

21. Tergum 1 and anterior part of tergum 2 largely ferruginous; free margin of the 

clypeal process rounded ^vyomingensis Scullen 

Black with creamy white or yellow markings only 22 

22. Markings yellow 23 

Markings creamy white 25 



344 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii6 

23. Band on tergum 2 broader than on other terga; clypeal process emarginate 

(Eastern States, rarely west to North and South Dakota, Nebraska, and 

Kansas) atramontensis Banks 

Bands on terga 2 to 5 subequal in width 24 

24. Apex of clypeal process rounded (California and bordering states) . 

aequalis aeqiialis Provancher 

Apex of clypeal process somewhat truncate and emarginate (California and 

bordering areas) varians Mickel 

25. Apical end of the clypeal process acute or rounded, not emarginate (Idaho, 

adjoining areas, and mountains of central New Mexico). 

aequalis idahoensis Scullen 
Apical end of the clypeal process slightly emarginate 261 

26. Apical denticle on the mandible the larger; lateral apices of the clypea 

process not extended (Rocky Mountains, west to eastern Washington' 
east to North Dakota, north to Alberta, and south to northern Texas). 

calochorti Rohwer 
Medial denticle on the mandible the larger; lateral apices of the clypeal 
process extended to form a distinct denticle (southern California). 

macswaini Scullen 

27. Clypeal process sides distinctly converging to a smaller apex 28 

Clypeal process sides subparallel 31 

28. Markings showing considerable ferruginous or amber (eastern slope of the 

Rocky Mountains and adjoining areas) vicina Cresson 

Markings black and yellow or cream, no ferruginous or amber 29 

29. Markings black and white; apex of clypeal process rounded. 

aequalis bolingeri Scullen 
Markings black and creamy yellow; apex of clypeal process truncate . . 30 

30. Pygidium sides subparallel (Central and Northeastern States). 

astarte Banks 
Pygidium sides converging to a narrow apex . . clyjKjata dakotensis Banks 

31. Bands on terga 2 to 5 broad and not emarginate. morata Cresson (in part) 
Bands on terga 2 to 5 not all broad, some narrow or emarginate .... 32 

32. Bands on terga 2 to 5 subequal in width; tergum 1 with considerable fulvous. 

nebrascensis H. S. Smith 
Band on tergum 2 broader than those on terga 3 to 5 33 

33. Markings with considerable amber or shades of brown. 

morata Cresson (in part) 
Markings black and yellow, no amber or brown 34 

34. Band on tergum 2 deeply emarginate (Eastern States, rarely in the Central 

States) clypeata prominens Banks 

Band on tergum 2 with little or no emargination 35 

35. Bands on terga 3 to 5 narrow but unbroken and broad laterally; band on 

tergum 2 sometimes slightly emarginate (Eastern States, rarely west to 

the Great Plains) clypeata clypeata Dahlbom 

Bands on terga 3 to 5 absent or reduced to lateral spots; band on tergum 2 
not emarginate (Great Plains, rare in the East) . clypeata gnarina Banks 

36. Large species, 16 mm. or longer 37 

Small species, 12 mm. or less in length 42 

37. Clypeal process with a single acute apex verticalis F. Smith 

Clypeal process bicuspidate 38 



WASP GENUS CERCERIS — SCULLEN 345 

38. Lateral apices of the clypeal process considerably prolonged giving the 

appearance of horns or the points of a new moon 39 

Lateral apices of the clypeal process not greatly prolonged 40 

39. Terga of the abdomen almost entirely yellow . . . frontata frontata Say 
Terga of the abdomen with yellow bands much reduced. 

frontata raui Rohwer 

40. Distance between apices of the clj^peal process very much less than one- 

third the distance between the eyes; surface of the clypeal process not 

lunar shaped mimica Cresson 

Distance between apices of the clypeal process approximately equal to or 
greater than one-tliird the distance between the eyes; surface of the 

clypeal process somewhat lunar shaped 41 

4L Mostly black to dark amber with limited yellow. 

bicorniita bicornuta Guerin 
Mostly light amber and yellow . . bicornuta fidelis Viereck and Cockerell 

42. Tergum 2 black or dark amber without a yellow band (small evanescent 

yellow spots may appear) 43 

Tergum 2 with a yellow band 44 

43. Clypeal process as long as or longer than its width at the base and with a 

definite overhang; tergum 1 with a creamy yellow lateral border, black 

medially azteca Saussure 

Clypeal process very low without an overhang extension (somewhat similar 
to that of C. bicornuta Guerin) ; tergum 1 with a broad medial yellow 
patch and black lateral borders tolteca Saussure 

44. Clypeal process with a single acute apex (mostly Northeastern States, 

rarely in Texas) nitida Banks 

Clypeal process with two or more apices 45 

45. Clypeal medial lobe with a single medial acute denticle flanked by a pair of 

blunt denticles, all black melanthe Banks 

Clypeal medial lobe with a pair of yellow denticles connected by a low 
concave carina boharti ScuUen 

Group I 

L Elevated process on the free surface of the central clypeal lobe distinctly 

separated from the anterior border 2 

Clypeal process or denticles confined to or embodying the free anterior 
border 21 

2. No red area on the abdomen or thorax 3 

Distinct red area covering one or more anterior abdominal terga and possibly 

extending onto the propodeum of the thorax 15 

3. Elevation on the surface of the clypeus a single, low, cone-shaped process 

with a pair of erect denticles below near the clypeal border 4 

Elevation scoop shaped, truncate, or double pointed 5 

4. Black with yellow markings vanduzeei vanduzeei Banks 

Black with cream markings vanduzeei cburnea ScuUen 

5. Clypeal process extended so that the length is equal to or greater than the 

width; often inverted scoop shaped 6 

Clypeal process either short and broad or otherwise modified 8 

6. Apical border of clypeal process smooth and straight with rounded corners. 

truncata Cameron 

Apical border of clypeal process sinuate, giving the effect of three slight 

extensions 7 



346 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

7« Markings yellow kennicottii kennicottii Cresson 

Markings cream kennicottii beali Scullen 

8. Extension near the clypeal border emarginate. 

convergens Viereck and Cockerell 
Process not in the form of an emarginate lobe near the clypeal border ... 9 

9. Process broad and scoop shaped, much broader than long 10 

Process not scoop shaped; very low on the clypeus 12 

10. Mesosternal process absent poculum Scullen 

Mesosternal process present 11 

11. Clypeal process very broad with the apical border straight except for the 

extremities, which are curved downward; process cream except for black 

border chilopsidis Viereck and Cockerell 

Apical border of the clypeal process emarginate with the lateral angles ex- 
tended and rounded at the apex; process black except (in some specimens) 
for a small yellow spot in the center crandalli Scullen 

12. Clypeal process with its free margin converging to a blunt point, below which 

are two distinct denticles krombcini Scullen 

Clypeal process not converging to a point 13 

13. Center of face and clypeaus creamy white to white 14 

Center of face and clypeus black echo atrata Scullen 

14. Clypeal process subequal in width to the clypeal border extension ; pygidium 

pyriform acanthophila Cockerell 

Clypeal process about one-half as wide as the clypeal border extension; 
pygidium oval in form neahminax Scullen 

15. Clypeal elevation somewhat cone shaped with a single point that may be 

more or less rounded or blunt but not truncate 16 

Clypeal elevation scoop shaped, definitely truncate or otherwise modified . 17 

16. Clypeal process pointing ventrad toward the clypeal border; without erect 

denticles ventrad of the single elevation argia Mickel 

Cone-shaped process approximately vertical to the clypeal surface with two 
small erect black denticles ventrad of the cone-shaped process. 

conifrons Mickel 

17. Clypeal elevation a truncate cone 18 

Clypeal elevation not a truncate cone 19 

18. Silvery white markings; tegulae smooth and not elevated. 

crotonella Viereck and Cockerell 
Yellow markings; tegulae pitted and elevated irene Banks 

19. Clypeal process scoop shaped; face largely reddish yellow . . blakei Cresson 
Clypeal process not scoop shaped; face not reddish yellow 20 

20. Clypeal elevation reduced to little more than a short ridge parallel to the 

anterior clypeal border echo echo Mickel 

Clypeal elevation prominent and rounded with a deep incision on the anterior 
margin; single dark denticles on the lateral borders of the incision. 

bridwelli Scullen 

21. Distinct red area covering one or more anterior terga of the abdomen and in 

some cases extending on to the propodeum of the thorax 22 

No red area on the abdomen or thorax 23 

22. Propodeum largely black rufinoda rufinoda Cresson 

Propodeum red except enclosure, which is black. 

rufinoda crucis Viereck and Cockrell 

23. Clypeal border with a thin lobed medial process butleri Scullen 

Clypeal border with two widely separated hyaline points 24 



WASP GENUS CERCERIS — SCULLEN 347 

24. Markings silvery white; tegula smooth, not elevated. 

finitima vierecki Banks 
Markings yellow; tegula elevated and pitted 25 

25. Bands on terga narrow and deeply emarginate; propodeum black, immacu- 

late; occiput black except for small yellow spots back of the compound 

eyes finitima finitima Cresson 

Yellow markings greatly extended; yellow bands on terga wide with little 
or no emargination; considerable yellow on the propodeum and occiput. 

finitima citrina Scullen 

Group II 

1. Enclosure surface deeply ridged at 45" angle to base 2 

Enclosure surface not ridged 4 

2. Pygidium with sides converging uniformly to a narrow apical point; length 

20 mm 3 

Pygidium pyriform with sides concave on the apical half; length 14 to 15 mm. 

completa Banks 

3. Yellow with very limited amber markings, very little or no black. 

grandis grandis Banks 
Black with yellow markings grandis percna Scullen 

4. Abdomen black with one complete, wide ivory band on second tergum. 

fumipennis Say 

Abdomen largely yellow, light amber, or with light bands on more than one 

tergum 5 

5. Posterior half of abdomen (from center of third tergum) brownish yellow; 

thorax black except for broken band on pronotum and metanotum . . 6 

Thorax largely black or amber yellow; yellow bands on all terga or abdomen 

all yellow 7 

6. First and second abdominal segments black . . dilatata dilatata Spinola 
First abdominal segment reddish, second tergum with a limited amber yellow 

patch and red shading dilatata chisosensis Scullen 

7. Black with yellow markings, little or no amber. 

californica californica Cresson 

Head and thorax light amber marked with yellow; abdomen largely yellow 

or with some light amber californica arno Banks 

Group III 

1. Clypeal process broad and lunar or crescent shaped with the free margin 

subequal to the process in width 2 

Clypeal process narrowing to a margin apically subequal in width to one- 
half the width of the process 7 

2. Process with an undivided short but continuous lamella extending over 

nearly the entire margin of the process mandibularis Patton 

Lamella of process divided into two distinct oval lobes 3 

3. Process equal to approximately two-thirds of the distance between the 

eyes 4 

Process equal to about seven-eighths of the distance between the eyes . . 6 

4. Amber and yellow markings cochisi Scullen 

Black and yellow markings * 5 



348 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

5. Abdomen almost entirely yellow, occasionally some black on the first 

tergum ; prominent yellow patch on propodeum often C-shaped. 

sandiegensis ScuUen 

Most terga deeply emarginate with black; band on tergum 2 broader than 

others; propodeum with little or no yellow . compacta compacta Cresson 

6. Black with limited yellow markings; wide yellow band on distal half of 

tergum 2; no red or amber . . . flavofasciata flavofasciata H. S. Smith 

Head and thorax mostly reddish amber; first tergum reddish amber; second 

tergum reddish amber anteriorly, often wath a yellow line posteriorly; 

terga 4, 5, and 6 dark amber to black . flavofasciata floridensis Banks 

7. Lamella bent at about 45° angle to plane of clypeal process. 

squamulifera Mickel 
Lamella slightly or not at all bent from the plane of the clypeal process . . 8 

8. Lamella undivided to base, border might be slightly emarginate .... 9 
Lamella divided to base, divisions more or less completely separated . . 10 

9. Black and yellow; no reddish amber robertsonii robertsonii Fox 

Prothorax, scutellum, metanotum, basal two or more terga, and legs mostly 

reddish amber robertsonii emmiltosiis Scullen 

10. Black and yellow robertsonii bifidus Scullen 

Prothorax, scutellum, metanotum, basal two terga, and legs reddish amber 
with traces of yellow robertsonii miltosus Scullen 

Group IV 

1. Length 1.5 to 18 mm.; punctation deep and close. 

macrosticta Viereck and Cockerell 
Length 10 to 12 mm.; punctation fine and more widely separated .... 2 

2. Body colors black and yellow with very little or no amber. 

femurrubrum athene Banks 
Body colors amber and cream with very little black or none at all. 

femurrubrum femvirrubrum Viereck and Cockerell 

Group V 

1. Clypeal elevation low and rounded without a distinct point 2 

Clypeal elevation not low and rounded but sides converging to a distinct 

apex, which may be blunt, approximately perpendicular to the surface; 
lower surface of elevation smooth and concave or fiat 5 

2. Color markings dark red rufa ScuUen 

Color markings yellow or cream 3 

3. Subequal narrow bands on all terga compar compar Cresson 

Band on tergum 2 absent or reduced to small lateral patches on the posterior 

margin 4 

4. Usually a single elongate patch and rarely a second smaller yellow spot on 

the propodeum; band on tergum 3 more or less emarginate on the anterior 
margin; small yellow spots usually on the lateral posterior angles of ter- 

g'lm 2 compar geniculata Cameron 

Markings on propodeum fused, or nearly so, to form a hook-shaped patch; 
wide complete band on tergum 3; somewhat smaller than the latter sub- 
species compar orestes Banks 

5. Second tergum with a band equal to or wider than that on other terga. 

zelica Banks 

Second tergum immaculate or with small lateral spots on the posterior margin; 

bands on other terga subequal 6 



WASP GENUS CERCERIS — SCULLEN 349 

6. Markings all creamy white; band on third tergum, others a narrow line (South- 

west desert area, New Mexico) insolita albida ScuUen 

Markings on body yellow 7 

7. Single yellow patch on the propodeum; band on tergum 3 emarginate; py- 

gidium fuscous; hindfemora variable shades of amber (East Central 

States) insolita insolita Cresson 

A small secondary yellow spot often appearing on the propodeum ; band on 
tergum 3 broad and showing little or no cmargination ; pygidium ferru- 
ginous; hindfemora black with a creamy j-ellow patch near the apical end 
(Southwest desert area, Arizona) insolita atrafeniori ScuUen 

MALES 

1. Stigma very dark to black 2 

Stigma light amber 3 

2. Length not over 9 mm. ; complete bands on one or more terga in addition to 

tergum 2 Group I 

Length 10 to 11 mm.; complete broad band on posterior half of tergum 2; 
other terga with lateral markings or broken lines only, very rarely solid 
lines fumipennis Say (of Group II) 

3. Width of clypeal hair lobes greater than the distance between them and 

extending onto the medial clypeal lobe 4 

Width of the clypeal hair lobes no greater than the distance between them 
and not extending onto the medial clypeal lobe 7 

4. Hair lobes meet medially or nearly so; tergum 2 immaculate; posterior half 

of tergum 3 and all of terga 4, 5, and 6 fulvous .... azteca Saussure 

Hair lobes distinctly separated medially; bands on terga 4, 5, and 6 not 

all broad, at least some are narrow or deeply emarginate 5 

5. Medial denticle of the clypeal margin bicuspidate Group IV 

Medial denticle of the clypeal margin not bicuspidate 6 

6. Pygidium semioval; lateral clypeal lobe heavily setose . . tolteca Saussure 
Pygidium elongate, approximately twice as long as it is broad; lateral 

clypeal lobe very lightly setose Group V 

7. Medial lobe of clypeus less than half as wide as lateral lobe . . . Group III 
Medial lobe of clypeus subequal to or wider than lateral lobe 8 

8. Prominent cluster of bristles at the side of the pygidium 9 

Bristles adjacent to pygidium sparse 12 

9. Bands on all terga narrowly interrupted along midline 10 

Not all bands on terga narrowly interrupted along midline 11 

10. Pygidium broad, width subequal to length; pubescence normal. 

verticalis F. Smith 

Pygidium much longer than broad; pubescence very long on most body 

parts; pubescence on venter very prominent mimica Cresson 

11. Background color largely black bicornnta bicornuta Guerin 

Background color largely ferruginous (Southwest desert area) . 

bicornnta fidelis Viereck and Cockerell 

12. Basal width of second tergum subequal to one-third the greatest width of 

the segment Group II 

Basal width of second tergum subequal to or greater in width than one-half 
of the greatest width of the segment 13 

13. More or less ferruginous or amber replacing the darker color, especially along 

the margins of the dorsal abdominal markings 14 

Black with yellow, cream-colored, or white markings only 16 

742^63—65^—2 



350 PROCEEDINGS OF THE NATIONAL MUSEUM tol. ne 

14. Pygidium with sides diverging slightly apically and convex on the apical 

end niorata Cresson 

(also vicina Cresson) ^ 

Pygidium with sides converging slightly and the apical end slightly 

concave 15 

15. Bands of yellow on the tcrga with little or no emargination. 

frontata frontata Say 
Bands on the terga deeply emarginate anteriorly . . frontata rani Rohwer 

16. Band on tergum 2 distinctly wider than the bands on other terga . . 17 
Bands on all terga subequal 19 

17. Small species, approximately 8 mm. in length nitida Banks 

Medium in length, 10 to 12 mm 18 

18. Band on tergum 2 deeply emarginate .... clypeata prominens Banks 

(probablj- also halone Banks) 
Band on tergum 2 not emarginate or, if so, very little. 

clypeata clypeata Dahlbom 
(probably also clypeata gnarina Banks, dakotensis Banks, posticata 
Banks,* atraviontensis Banks, rhois Rohwer*) 

19. Small species about 9 mm. in length; clypeal denticles reduced to two 

(Northeastern United States) astarte Banks 

Medium to large species, 10 mm. or longer ; tlu-ee more or less distinct clypeal 
denticles 20 

20. Denticles on the clypeal margin separated bj^ a distance subequal to one- 

fifth the width of the medial lobe; a distinct transverse ridge just above 

the denticles (Eastern States) deserta Say 

Denticles on the clypeal margin in approximate conjunction; no transverse 
ridge on the surface of the medial lobe 21 

21. Markings yellow 22 

Markings white to cream 26 

22. Lateral clypeal denticle prominent; the medial clypeal denticle greatly 

reduced or absent 23 

Three clypeal denticles subequal 24 

23. Medial clypeal lobe with a distinct transverse swelling about the level of the 

anterior tentorial pits (Central States south to Texas). 

occipitomaculata Packard 

Medial clypeal lobe ^vithout a distinct transverse swelling but generally 

convex nigrescens arelate Banks 

24. Length about 10 mm.; denticles on the clypeal border very indistinct (Cali- 

fornia and adjoining states) nigrescens munda Mickel 

(probably also aequalis aequalis Provancher, varians Mickel, dentifrons 
Cresson) 
Length about 13 mm. ; denticles on clypeal border more distinct 25 

25. Seventh tergum with yellow to the sides of the pygidium (West Coast 

States) tepaneca Saussure 

Seventh tergum immaculate (Central Plains States) sexta Say 

(also posticata Banks, rhois Rohwer) 



* The males of morata Cresson and vicina Cresson so far are inseparable except 
by association with the females. 

* The type of posticata Banks runs to couplet 18b. It is a unique that may be 
sexta Say with an abnormally wide band on tergum 2. The type of rhois Rohwer, 
which also runs to this couplet, could be the male of clypeata gnarina Banks, but 
the type locality is out of the known range of that species. 



WASP GENUS CERCERIS — SCULLEN 351 

26. Length 14 to 15 mm. (North Central States) stigmosalis Banks 

Length 9 to 10 mm 27 

27. Propodeum immaculate (Northern States from coast to coast). 

nigrescens nigrescens F. Smith 
Propodeum with large white patches 28 

28. Enclosure smooth (eastern slope of the Rocky Mountains). 

calochorti Rohwer 
Enclosure more or less coarsely ridged (southern California) . 

macswaiiii Scullen 

Group I 

1. Showing shades of ferruginous or red on the first abdominal tergum and 

bordering areas 2 

Colors black with yellow, white, or cream markings only, no ferruginous or 
red markings 6 

2. Markings creamy white or white 3 

Markings yellow 5 

3. Terga 2, 3, and 4 with broad bands of creamy white, in some specimens 

the fourth showing slight emargination; punctation sparse (known only 

from southern California and southern Arizona) . . . bridwelli Scullen 

Wide bands on terga 2 and 3 only, or aU bands subequal and of medium 

width; punctation crowded or average 4 

4. Bands on terga 2 and 3 broad medially but narrowing laterally, bands on 4 

and 5 narrow; punctation medium (known only from southwestern Arizona 

and southern Texas) crotonella Viereck and Cockerell 

Bands on terga 2 to 6 subequal and of medium width; punctation crowded 
(general over western arid sections but more common in the Southwest). 

conifrons Mickel 

5. Band on tergum 2 with a deep V-shaped emargination, bands on terga 3 to 

6 broad but deeply emarginate and often completely divided (extreme 

Southeastern States) blakei Cresson 

Band on tergum 2 broad and with little or no emargination, bands on terga 

3 to 6 of medium width with broad and moderately deep emarginations 

(recorded from most states except the New England States, the Great 

Lake Area and the extreme Southeast) . . . rufinoda rufinoda Cresson 

(also echo echo Mickel ;8 argia Mickel and irene Banks, when recognized, 

probably will run to this couplet also) 

6. Markings on tlie abdomen creamy 7 

Markings on the abdomen yellow 12 

7. Propodeum with medium patches of creamy white; usually two small spots 

near the apex of the enclosure; first tergum cream except for a black patch 
at the base (southern Arizona, New Mexico, and Texas). 

truncata Cameron 

Propodeum and enclosure immaculate; tergum 1 either immaculate or with 

only a small cream patch on the dorsum 8 



8 The males of rufinoda rufinoda Cresson and echo echo Mickel so far are indis- 
tinguishable except as they are associated with the females. Some specimens of 
males taken at the same time and place as some females of echo echo Mickel are 
more cream than the males usually accepted as the males of rufinoda rufinoda 
Cresson. These forms with lighter markings may prove to be the males of echo 
echo Mickel, 



352 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

8. Baud on tergum 3 missing, a broken line, or divided into three sections; spot 

on first tergum very small or absent; lateral lobes of the clypeus black . 9 
Baud on tergum 3 unbroken but emarginate 10 

9. Band on tergum 3 divided at two points to form three parts subequal in 

length (in some specimens these divisions are represented by indentations 
only); entire face black except for two large frontal eye patches; medial 
denticle on the clypeal margin more prominent than the others (west of 
the Rocky Mountains from British Columbia to Texas). 

convergens Viereck and Cockerell (in part) 
Band on tergum 3 usually a broken or evanescent line; lateral lobes and 
lower margin of the medial lobe of the clypeus black; frontal area black 
with a small evanescent spot of cream (Southwestern States). 

kennicottii beali ScuUen 

10. Tergum 1 immaculate; apex of median lobe and lateral lobes of clypeus 

black; face otherwise creamy yellow cleome Rohwer 

Tergum 1 with a distinct patch 11 

11. Clypeal denticles clearly defined; tegulae showing a tendency to be elevated 

and inflated; light apical end of the third femora demarked somewhat 

obliquely (Southwestern States) finitima vierecki Banks 

Clypeal denticles less elevated and distinct ; tegulae not elevated or inflated ; 
light apical end of the third femora demarked nearly at right angle to the 
femur 12 

12. Scutellum immaculate; pygidium nearly as broad as it is long. 

crandelli ScuUen 
Scutellum ■with two cream patches; pygidium about one and one-half as 

long as it is broad acanthophila Cockerell 

(also carrizoensis Banks, cockerelli Viereck) 

13. Tegulae elevated 14 

Tegulae not elevated 16 

14. Tegulae smooth; enclosure smooth except for a medial groove and a limited 

number of deep pits along the lateral margins (southern Arizona). 

krombeini Scullen 

Tegulae pitted; enclosure heavily regose or ridged subparallel to the anterior 

border 15 

15. Band on tergum 3 deeply emarginate; no yellow marks on the genae or 

occiput (more common in the Eastern States but ranges into California). 

finitima finitima Cresson ' 
Broad band on terga 2, 3, and often 4 with little or no emargination ; com- 
monly large yellow patches back of the eyes and a broken band on the 
occiput (southern and central California) . . . finitima citrina Scullen 

16. Band on tergum 3 divided at two points to form three parts subequal in 

length (in some specimens these divisions are represented by indentations 
only); entire face black except for two large frontal eye patches; medial 
denticle on the clypeal margin more prominent than the others (west of 
the Rocky Mountains from British Columbia to Texas). 

convergens Viereck and Cockerell (in part) 
Band on tergum 3 reduced to a broken evanescent line or completely elimi- 
nated medially; lateral lobes of the clypeus black; apical border of the 
medial lobe of the clypeus black; frons black (recorded from most of the 
Eastern States and Southwest) .... kennicottii kennicottii Cresson 



^ C. arizonella Banks appears to be a form between C. f. finitima Cresson and 
C. f. citrina Scullen. 



WASP GENUS CERCERIS — SCULLEN 353 

Group II 

1. Tergum 2 black or with more or less ferruginous 2 

Tergum 2 all yellow or with a broad yellow band 3 

2. Terga 1 and 2 black, basal half of terguna 3 black; posterior half of tergum 

3 and reiBaining posterior terga brownish yellow (Arizona, New Mexico, 

and southern Texas) dilatata dilatata Spinola 

Black of basal abdominal segments more or less replaced with red (Big Bend 
National Park and Davis Mts. in Texas, southern New Mexico, and 
southern Arizona) dilatata chisosensis ScuUen 

3. All terga of the abdomen yellow or very light fulvous except for a trace of 

emargination at the base 4 

Bands on terga 3 to 6 deeply emarginate anteriorly in middle 6 

4. Thorax black to dark fuscous background; first abdominal segment may show 

more or less red californica californica Cresson (in part) 

Thoracic background fulvous 5 

5. About 10 mm. in length; enclosure lightly ridged (southern California and 

adjoining areas californica arno Banks 

About 14 mm. in length; enclosure relatively smooth but with a few pits 
in the lateral angles (southern California and adjoining areas) . 

grandis grandis Banks 

6. Length about 10 mm.; legs amber and yellow (this species shows great varia- 

tion in amounts and shades of yellow and red throughout its range west 
of the Rocky Mountains from British Columbia to Texas). 

californica californica Cresson (in part) 

Length about 12 to 15 mm.; legs black and yellow; emarginations on terga 

usually narrow but deep, that on tergum 2 with a tendency to be enclosed 

by yellow (California and southwestern Oregon) . . . completa Banks 

Group III 

1. Broad bands with little or no emargination on terga 3 to 6 2 

Bands on terga 3 to 6 either greatly reduced or with deep emarginations . 3 

2. Background black or dark fuscous (southern California). 

sandiegensis Scullen 
Backgound ferruginous (southern Arizona and adjoining areas). 

cochisi Scullen 

3. Black with yellow and/or fulvous markings, no red or ferruginous ... 5 
Black with red and/or ferruginous markings (Florida and adjoining areas) . 4 

4. Marginal extension of the clypeus about as wide as half the length of the 

epistomal suture; mandibular denticles strong; enclosure smooth. 

flavofasciata floridensis Banks 
Marginal extension of the clypeus about equal in width to the length of the 
epistomal suture; mandibular denticles small; enclosure ridged. 

rufopicta F. Smith* 

5. Lower part of the clypeus showing considerable black and the yellow area 

of the medial lobe tapering apically to an acute point; metanotum yellow 
(common through the Eastern States from southern Minnesota, southern 
Wisconsin, southern Michigan, and New England south to the Gulf of Mex- 
ico and west to Utah and the Southwest) . compacta conipacta Cresson 



8 Field observations may prove C. rufopicta F. Smith to be the male of C. 
robertsonii emmiltosus Scullen or C. roJtertsonii miltosus Scullen. 



354 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Face yellow with little or no black margin on the clypeus; yellow of face does 
not end apically in an acute point; metanotum immaculate or with very 

little yellow 6 

6. Bands on terga 3 to 6 reduced to an evanescent line or to small lateral patches 
(scattering through the East from Kansas and Nebraska to the Atlantic 

States) flavofasciata flavofasciata H. S. Smith 

Bands on terga solid but relatively narrow lines (distribution similar to the 

latter species) robertsonii robertsonii Fox 

(probably robertsonii hifidus Scullen, mandihularis Patton) 

Group IV 

1. Length about 15 mm.; markings dark fuscous, ferruginous, and yellow. 

macrosticta Vireck and Cockerell 
Length about 11 mm.; markings black and creamy white. 

feniurrubrum Viereck and Cockerell 

Group V 

1. Markings black with rufous (Florida) rufa Scullen 

Markings black and yellow or creamy white 2 

2. Tergum 2 immaculate or with small lateral spots only 3 

Tergum 2 with a solid but narrow band (general over Eastern States from 

Nebraska, Kansas, and Texas to the Atlantic Coast). 

compar conipar Cresson 

3. Without a mesosternal process or ridge (southern Arizona). 

compar orestes Banks 

(Probably zelica Banks) 

With a mesosternal process or ridge (sometimes very slight) 4 

4. Mesosternal process broad and considerably extended; markings yellow 

(only one recorded from southern Arizona), compar geniculata Cameron 
Mesosternal process reduced to a slight ridge 5 

5. Markings yellow (Eastern States from the Missouri River and Texas to 

the Atlantic States) insolita insolita Cresson 

Markings creamy white (southern Arizona, New Mexico, and southwestern 
Texas) insolita albida Scullen 

(Additional male belonging to Group V but not yet known: 
insolita atrafemori Scullen) 

Group 1 

This group ^ is distiuguished by the following characters: (1) small 
size, (2) very dark stigma, (3) punctation usually very crowded, (4) 
anterior abdominal segments and propodeum often more or less red- 
dish, (5) mandibles of female with two denticles, (6) mesosternal 
tubercle on the female, (7) small and widely separated hair lobes on 
the male, (8) apical segment of the antennae normal in the male. 



8 This group is the one Banks (1947, p. 26) calls Apiraptrix (Shestakov). 
Banks misspelled this "Apiratrix." In a personal letter, de Beaumont informs 
the present writer that this group is not Apiraptrix (Shestakov). The latter is 
related to C. rybyensis (Linnaeus) Schletterer. 



WASP GENUS CERCERIS — SCULLEN 355 

1. Cerceris acanthophila Cockerel! 

Figures 1, 108a,b,c 

Cerceris acanthophila Cockerell, 1897, p. 135. — Ashmead, 1899, p. 296. — Viereck 
and Cockerell, 1904, p. 138.— Viereck, 1906b, p. 234.— Banks, 1947, p. 30.— 
Scullen, 1951, p. 1004; 1960, p. 75. 

Cerceris minax Mickel, 1917a, p. 339. — Banks, 1947, p. 34. — Scullen, 1951, p. 
1009. 

Cerceris huachuca Banks, 1947, p. 29. — Scullen, 1951, p. 1008. 

Female. — ^Length 8 to 9 mm. Black with yellow to creamy-white 
markings, normally pitted, clothed with short silvery hairs. 

Head sUghtly wider than the thorax; front creamy white below 
antennae except apical borders, which are dark; vertex, occiput, and 
genae black except for small spots back of compound eyes, which are 
creamy white; clypeal border sHghtly sinuate; cl3T)eal elevation close 
to the anterior border, very short and subequal in width to the exten- 
sion of the clypeal border, shghtly emarginate; mandibles with two 
separated, centrally located subequal denticles; yellow at the base 
becoming dark amber apically; antennae normal in form, creamy- white 
patch on the scape, pedicel dark amber, flageUum light amber below, 
dark amber above. 

Thorax black except for two elongate patches on the pronotum, the 
metanotum, and the tegulae, which are creamy white; propodeum im- 
maculate, heavily and closely pitted except for the enclosure, which is 
moderately rugose; mesosternal tubercle small and black; legs black 
except for apical ends of femora, fore- and midtibiae, fore- and mid- 
tarsi, basal two-thirds of the hindtibiae, and basal third of the hind- 
tarsi, which are hght yellow; wings subhyaline, stigma medium dark 
(hghter than is usual in the group). 

Abdomen: terga 1 to 5 black with creamy-white bands deeply 
emarginate on 2 to 5; venter black except lateral yellow spots on 
sternites 3 and 4 (immaculate in some specimens) ; pygidium pyrif orm, 
widest near the base, narrowed apically with a cluster of small bristles 
on each side inserted on the apical sternite. 

Male. — Length 7 nmi. Black with creamy-white markings; 
punctation and pubescence average. 

Head shghtly wider than the thorax; black except for the entire 
face, basal part of mandibles, and the scape, all of which are creamy 
white; peduncle and flagellum ferruginous above and hght fulvous 
below; three cljrpeal border denticles distinct but not prominent; hair 
lobes narrow; mandibles smooth with a shght medial elevation. 

Thorax black except for a divided band on the pronotum, the 
scutellum, the metanotum, a small spot on the pleuron, and a patch 
on the tegulae, all of which are creamy white; mesosternal tubercle 



356 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

absent; legs black basally to the apical ends of all femora; fore- and 
midlegs are creamy white apicaUy except the tarsi, which become light 
fulvous ; the liindlegs are creamy white on the apical end of the femora 
and basal half of the tibiae, beyond which they are largely black ; wings 
subhyahne but sHghtly clouded apically. 

Abdomen black except for a small patch on tergum 1, broad band 
on tergum 2, narrower and emarginate bands on terga 3, 4, and 5 and 
a small spot on tergum 6, all of which are creamy white; venter im- 
maculate ; p3^gidium mth sides sUghtly convex, ends subequal in width 
and broadly rounded. 

Many of the specimens show the markings more yellow than white. 
The hghter forms are more common in the Pacific Northwest and in 
the southv/estem desert area. It is possible subspecies should be 
recognized. The extent of the Hght markings varies. In some speci- 
mens spots appear below the wing attachments and on the meso- 
sternal tubercles. 

The cljrpeal process easily distinguishes the female of acanthophila 
CockereU from closely related species such asjinitima Cresson. 

Types. — A male of C. acanthophila Cockerell is in the Philadelphia 
Academy of Natural History, ANSP type no. 10038, from Deming, 
N. Mex. This is considered the lectotype designated by Cresson in 
1928. Another male, apparently from the same series, is in the 
National Museum, Washington, D.C., type no. 3409. A second 
specimen with a blue ANSP paratype label 10038 is also in the Phila- 
delphia Academy of Natural History. 

The holotype female and allotype male of C. minax Mickel from 
Sacramento, CaUf., are at the University of Nebraska. Three para- 
type females from Auburn, Calif., are also at the University of 
Nebraska. 

The holotype male of C. huachuca Banks from Patagonia, Ariz., is 
in the Museum of Comparative Zoology, Harvard University, type 
no. 27636. 

Distribution. — West of the Continental Divide from British 
Columbia into Mexico. 

Prey record. — None. 

Plant record. — Achillea (Idaho, Oregon), Achillea millefolium 
(Oregon), Asclepias mexicana (Oregon), Boerhaavia erecta (New 
Mexico), Daucus carota (CaHfornia, Idaho, Oregon), Eriogonum 
(Cahfornia, Idaho, Oregon)^ Eriogonum fasciculatum var. polifolium 
(California), Foeniculum vulgare (California), Harmizonia fasciculatum 
(Cahfornia), Helianthus (Idaho), Melilotus (Cahfornia), Melilotus 
alba (Oregon), Pastinaca sativa (Utah), Rhamnus (California), Salsola 
kali tenuifolia (Utah), Solidago (New Mexico, Oregon), Tamerix 
(Utah), Zizyphus lycioides (New Mexico). 



WASP GENUS CERCERIS — SCULLEN 



357 




rN- 


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[ 


^/ ' T 


* "r - 


-t^y^ 


; 


;^~i. 


v^ ~, 


/ 

I -^ ' 


N 


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Figures 1-3. — Localities of: 1, C. acanthophila Cockerell; 2, C. argia Micliel (females only); 
3, C. arizondla Banks. 



2. Cerceris argia Mickel 

Figures 2, 109a,b,c 
Cerceris argia Mickel, 1916, p. 412; 1917b, p. 453.— Scullen, 1951, p. 1005. 

Female. — Length 9 to 10 mm. Black with yellow markings except 
the propodeum and the basal end of the abdomen, which are reddish 
amber; more closely and finely pitted than most species; clothed with 
short silvery hairs. 

Head about one-sixth wider than the thorax, normally pitted, clothed 
with very short silvery hairs; front yellow below the antennae except 
for the anterior third of the clypeus, which is amber; occiput immacu- 
late, black in the type (many specimens show two converging, elongate 
yellow patches); genae black with a yellow spot back of each com- 
pound eye; clypeal border irregularly sinuate; clypeal process low 
with a single rounded end pointing ventrad, yellow with the free 
border amber; mandibles with two separated, centrally located sub- 
equal denticles, yellow at the base, amber apically; antennae normal in 
form, yellow area on scape, peduncle amber, flagellum hght amber 
below, dark amber above. 

Thorax closely pitted except on the scutellum, the metanotura, 
and the enclosure, clothed with short silvery hairs, black except for 
two elongate patches on the pronotum, two oval patches on the 
scutellum, the metanotum, a patch on the pleura, and the tegulae, all 
of which are light yellow; tegulae elevated and hghtly pitted; the 
propodeum is reddish amber except for the enclosure, which is black ; 
enclosure is smooth except for a mesal groove and lateral ridges, 
which are set at about a 45° angle to the mesal groove; mesosternal 
tubercles small and black; legs largely light amber except for yellow 



358 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

areas on apical ends of the forefemora, the entire fore tibiae, foretarsi, 
apical ends of the midfemora, basal ends of midtibiae, and traces of 
yellow on apical segments of the hindlegs; wings subhyaline, stigma 
lighter than normal for the group. 

Abdomen finely and closely pitted; first tergum and anterior half 
of the second tergum reddish amber; broad yellow bands on terga 2 
to 5 deeply emarginate ; anterior halves of terga 3 to 5 black ; sternum 
hght amber, becoming darker apically; pygidium broad at the base, 
gradually narrowing apically to a rounded end; small clusters of 
bristles on each side inserted on the apical sternite. 

Male. — Length 8 mm. Black with Ught yellow and dark ferru- 
ginous markings; punctation crowded; pubescence very short. 

Head subequal in width to the thorax; black except for the large 
frontal eye patches, most of the medial lobe of the clypeus, and the 
frons, all of which are light yellow; lateral lobes of the clypeus and the 
free border of the medial lobe of the clypeus are black; medial clypeal 
lobe \vith a small medial denticle and two less distinct lateral denticles ; 
hair lobes extending onto the medial lobe; mandibles without denticles ; 
antennae normal in form with a small yellow patch on the scape, the 
peduncle dark and the flagellum fulvous below becoming darker above. 

Thorax black except for a divided band on the pronotum, most of 
the tegulae, and a small spot on the pleuron below the wing, all of 
which are light yellow; tegulae slightly elevated and with scattered 
pits; enclosure deeply and coarsely pitted; mesosternal tubercles 
absent; legs black except the apical ends of all femora and the more 
apical segments, which are yellow infused with darker areas; the 
third trochanter and the adjoining end of the femur dark ferruginous; 
wings subhyaline, somewhat clouded at the apex and the stigma dark. 

Abdomen black except the first segment and the anterior margin 
of the second sternite, which are dark ferruginous, and a broad convex 
band on tergum 2, narrower emarginate bands on terga 3, 4, and 5, 
and a small patch on tergum 6, all of which are light yellow; venter 
all black except sternite 1 and the anterior border of sternite 2, which 
are dark ferruginous; pygidium deeply pitted, strongly convex 
laterally, proximal end rounded and distal end truncate. 

A study of many specimens from the same area shows the elongate 
yellow spots on the vertex are common and vary in size. The tegulae 
may also vary in elevation and smoothness. The female of C. argia 
Mickel resembles others of the rufinoda group but is distinguished 
easily by the clypeal process. 

Type. — The type female of C. argia Mickel is at the University of 
Nebraska. 



WASP GENUS CERCERIS — SCULLEN 359 

Distribution. — ^From Kansas and eastern Colorado southwest 
through Texas, Louisiana, New Mexico, and Arizona into Mexico. 

Prey record, — -None. 

Plant record. — Euphorbia marginata (Texas), Lepidium 
(Arizona), Solidago sp. (Nebraska). 

3. Cerceris arizonella Banks 

Figure 3 
Cerceris arizonella Banks, 1947, pp. 32-33. — ScuUen, 1951, p. 1005. 

Male. — 'Length 4 mm. Black with yellow markings except for 
the face, which is cream colored. Band on tergum 2 somewhat 
wider than the others. 

This species is known only from the type, which is close to finitima 
Cresson. It may prove to be an abnormally small specimen of that 
species. Until more positive characters for separating males are 
found, the author will let it remain a vaUd species. 

Type. — The type male, from Tempe, Ariz., is at the Museum of 
Comparative Zoology, no. 23538. 

Prey record. — 'None. 

Plant record. — -None. 

4. Cerceris blakei Cresson 

Figures 4, 110a,b,c 

Cerceris blakei Cresson, 1865, p. 121. — Packard, 1866, p. 61. — Cresson, 1887, 
p. 282.— DaUa Torre, 1897, p. 454.— Bridwell, 1898, p. 209.— Ashmead, 
1899, p. 295.— Banks, 1912a, p. 26.— ScuUen, 1951, p. 1005.— Krombein, 
1952c, p. 336; 1954b, p. 235; 1955, p. 234.— ScuUen, 1961, p. 45.io 

Cerceris elegans F. Smith, 1856, p. 467. — Cresson, 1865, p. 131. — Packard, 1866, 
p. 64.— Cresson, 1887, p. 282.— DaUa Torre, 1890, p. 200.— Ashmead, 1899, 
p. 295. 

Cerceris blackii [sic] Schletterer, 1887, p. 487. 

Female. — Length 9 to 10 mm. Black with reddish-amber mark- 
ings, the amber markings becoming yellow centrally in certain areas; 
deeply and closely pitted; clothed with very short silvery hairs. 

Head about one-thu'd wider than the thorax, black except sides of 
face, front, dorsal surface of the clypeal process, and small spots 
back of the compound eyes, all of which are yellow, and the clypeus 
below the process, which is amber; clypeal border sinuate, depressed 
medially; clypeal process scoop shaped with lateral sides subparallel 
and apical border smooth and uniformally curved ; mandibles with two 



1" Cerceris elegantissima Schletterer, 1887, was a new name proposed for 
Eucerceris elegans Cresson and not for C. elegans F. Smith as stated by ScuUen 
(1961, p. 45). 



360 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iis 

separated, centrally located denticles, the apical one much the larger; 
antennae normal in form, amber except the scape, which is yellow. 

Thorax black except two patches on the pronotum, two small spots 
on the propodeum, two small spots on the scutellmn, and the tegulae, 
all of which are yellow, and the metanotmn, which is yellow to amber; 
tegulae elevated and lightly pitted; enclosure smooth except for a 
longitudinal groove and pitted lateral borders; mesosternal tubercle 
small and black; legs yellow except for basal portions of femora, 
coxae, and trochanters, which are amber; wings uniformally clouded 
except for the stigma, which is dark a,mber. 

Abdomen: tergum 1 light amber; tergum 2 yellow, deeply emargi- 
nate with light amber; terga 3, 4, and 5 with anterior emarginations 
amber and deep, dividing, or ahnost dividing, the yellow into lateral 
triangular areas; venter light amber basally becoming dark apically; 
pygidium oval with the apical end truncate and the basal end 
rounded. 

The extent of the yellow markings varies. Yellow spots on the 
head and thorax may disappear completely in some specimens. 
Yellow and amber areas on the abdomen show much variation in 
their width and emargination. 

Male. — Length 8 mm. General coloration, punctation, and 
pubescence similar to the female. 

Head about one-third wider than the thorax; face yellow at the 
sides and below the antennae, remainder of head black; three amber 
denticles on the clypeal border, the medial one more broad and 
rounded than the lateral ones. 

Thorax black except for two patches on the pronotum, two spots on 
the scutellum, narrow baud on the metanotmn, and the tegulae, all 
of which are yellow; tegulae elevated and lightly pitted; enclosure 
deeply pitted to rugose; mesosternal tubercle absent; legs yellow 
except for basal two-thirds of femora, trochanters, and coxae, which 
are amber to black; antennae normal; wings uniformally clouded 
except for the stigma, which is dark amber. 

Abdomen: tergum 1 black with a broad divided amber band; 
terga 2 to 6 light amber to yellow, deeply emarginate anteriorly 
with black; venter dark amber, immaculate; pygidium slightly oval, 
truncate apically, rounded basally. 

The males are most apt to be confused with the males of jinitima 
Jinitima Cresson and kennicottii kennicottii Cresson. C. jinitima 
Jinitima Cresson lacks the amber shading. C. kennicottii kennicottii 
Cresson has the tegulae unelevated, smooth, and is black and yellow, 
without amber. 

Type. — The type female, from southwest Georgia (Charles A. 
Blake), is at the Philadelphia Academy of Natural Sciences, no. 1947. 



WASP GENUS CERCERIS — SCULLEN 



361 



Distribution. — Mostly in the Southeastern States from eastern 
Texas to North CaroUna and isolated records as follows: 

Illinois: Havans, Aug. 30, 1917; Meredosia, Aug. 22, 1898 (F.M.McE.) 
Aug. 1899. Kansas: Riley Co., July 17 (Popenoe). New Jersey: Camden Co. 
(Wm. J. Fox); Pemberton, Sept. 1, 1907. 

Prey records. — Derelomus basalis LeConte, Limnobaris confusa 
Bocheman, Anthonomus sexguttatus Dietz and Hyperodes sp. (all 
weevils). C. blakei Cresson seems to prefer the first species. Less 
commonly taken were the chrysomelid Graphops jioridana Blake and 
the tenebrionid Blakstinus interruptis (Say). All records are by 
Krombein (1963) from Florida. 

Plant records. — Aldenella tenuifolia (Florida, for nectar?), 
Asclepias (North Carolina), Bidens bipinnata (North Carolina), 
Borieria (Florida), Cyanotis (Alabama), Erigeron quercifolius (Florida), 
Eriogonum tomentosum (Florida), Koellia (North Carolina), lemon 
balm (Florida), Pinus serotina (North Carolina), Polygonum 
hyropiperoides (Florida), Quercus virginiana (North Carolina), Sabal 
eltonia (Florida, for prey), Satureia hortensis (Louisiana). 




Figures 4-6. — Localities of: 4, C. blakei Cresson; 5, C. bridwelli Sculler.; 6, C. buileri 
Scullen. 



5. Cerceris hridwelli, new species 

Figures 5, llla,b,c 

Female. — Length 8 mm. Black with creamy-yellow and amber 
markings; normally pitted, clothed with short silvery hairs. 

Head about 50% wider than the thorax; black except sides of 
face, compressed strip between the antennae, the clypeus, two round 
spots behind the compound eyes, proximal half of the mandibles, and 
the scape of the antennae, all of wliich are creamy yellow; clypeal 
border extended and sinuate; clypeal process prominent, rounded, and 



362 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

deeply depressed on the anterior side with the free border curved and 
ending on each side as a dark amber apex; mandibles with two sep- 
arated denticles, the apical one very much the larger; antennae normal 
in form, amber except the scape, which is creamy yellow. 

Thorax black except band on the pronotum, scutellum, metanotum, 
the tegulae, and two patches on the pleura, which are creamy yellow; 
tegulae smooth and not elevated; propodeum largely amber with a 
yellow patch on each side; enclosure black, lightly ridged at a 45° 
angle to the medial row of deep pits; mesosternal tubercle prominent 
and marked with creamy yellow; legs amber basally, changing to 
creamy yellow midway on the femora; wings subhyahne except the 
stigma, which is dark amber. 

Abdomen more sparsely but deeply pitted than the head and 
thorax; tergum 1 light amber with a partly divided yellow patch; 
terga 2 to 5 creamy yellow with a dark amber emargination on the 
anterior sides; pygidium oval with both ends rounded but narrowing 
apically; venter amber except two small spots on sternite 2 and 
narrow bands on sternites 3 and 4, which are creamy yellow. 

Male. — 'Length 7 mm. Black with creamy-white markings and 
limited ferruginous on the first two abdominal segments; punctation 
more widely spaced than average; pubescence short. 

Head black except for the face, minute spot back of the eye, base 
of the mandibles, and most of the scape, all of which are creamy 
white; clypeal border with three denticles on the medial lobe, the 
medial denticle more prominent than the lateral ones; surface of the 
medial lobe convex; hair lobes extend to the margin of the medial 
lobe of the clypeus; mandibles with slight evidence of a low, broad 
elevation but no distinct denticle; antennae with the apical segment 
slightly curved and slightly flattened on the apical end, fulvous 
beyond the peduncle, which is darker. 

Thorax black except for a divided band on the pronotum, two 
patches on the scutellum, the metanotum, a prominent patch on the 
pleuron, and the tegulae, all of which are creamy white; tegulae low 
and smooth; enclosure with a deeply pitted medial groove and deeply 
pitted lateral margins; mesosternal tubercles absent; legs dark fus- 
cous over the basal parts and largely creamy white over the apical 
ends of the femora and the more apical segments; the trochanter of 
the midleg shows a tendency to be lighter than the adjoining leg 
segments; wings subhyaline, stigma very dark. 

Abdomen black except the entire first segment is ferruginous and 
terga 2, 3, and 4 have broad bands, tergiun 5 has a narrow band and 
tergum 6 a narrow broken line, all of which are creamy white; venter 
with lateral creamy-white patches on sternites 2, 3, and 4 (that on 2 



WASP GENUS CERCERIS — SCULLEN 363 

very much smaller) ; p^^gidimn with sides convex and the apical end 
slightly smaller than the basal end. 

The male of bridwelli ScuUen is very close to the male of crotonella 
Viereck and Cockereil in size, sm-face sculptm^ing, and color pattern 
except there are three broad bands on the terga of the former and two 
broad bands on the terga of the latter; however, the band on torgum 
3 of the former occasionally shows a slight emargination or medial 
evanescent black spot. 

Yellow areas on both sexes show some variation in extent and smaller 
spots may disappear in some specimens; band on tergum 4 may be 
more or less emarginate. 

Types, — Type female and allotype male, from Imperial Co., Calif., 
June 1911 (J. C. Bridwell), are at the U.S. National Museum, no. 
66154. Paratypes are as follows: 

Arizona: 2 9 ?, 25 mi. W. Blythe, June 1937 (J. C. Elmore); 2d'c?, 11 mi. 
SW. Eloy, Pinal Co., June 19, 1953 (T. R. Haig) ; 9, Bd^cT, 10 mi. S. Toltec, 
Pinal Co., June 15, 21, 1953 (T. R. Haig); 9, Tucson, Apr. 25, 1940 (R. H. 
Crandall); 9, Yuma, May 4, 1918 (J. C. Bradley); 9, Yuma, May 6, 1939 
(M. R. Bohart). California: 9, Blythe, Riverside Co., May 8, 1947 (J. W. 
MacSwain, F. G. Linsley); 9, Borego, San Diego Co., Apr. 5, 1940 (R. M. 
Bohart); 5 9 9, lld^cT, Borego, San Diego Co., Apr. 25, 26, 27, 30, 1954, Croton 
californicus, Eriogonum inflalum (M. Wasbauer) ; cf, Borego, San Diego Co., 
May 2, 1952 (P. D. Kurd); 4 9 9 , 5cf cf, Borego Valley, San Diego Co., May 26, 
1954 (F. X. Williams); <?, Calexico, Aug. (Bequaert); 9, Colorado Desert, May 
12, 1935 (G. E. Bohart); 9, d", Desert Center, May 22, 1939 (E. P. Van Dyke); 
9, Dos PalmOS, Riverside Co., Apr. 3, 1934 (G. E. Bohart); 44 9 9, 57d'dr, 
Imperial Co., Experiment Farm, April, May, and June 1911, May and June 1912, 
Baccharis glutinosa (J. C. Bridwell); 7 9 9, 35crcf, 4 mi. E. Mecca, Riverside 
Co., July 16, 21, Aug. 15, 1956, Croton californicus (M. Wasbauer); 9 , San Felipe 
Creek, San Diego Co., July 8, 1933 (G. E. Bohart); 9, cf, Westmorland, May 
1932 (G. E. Bohart). Mexico: Baja California: 9, Los Animas Gulf, May 8, 
1921 (E. P. Van Duzee). 

Distribution. — Southern Arizona and southern California. 

Prey Record. — None. 

Plant Record. — As listed under paratypes. 

6. Cerceris butleri, new species 

Figures 6, 112a,b,c 

Female. — Length 7 mm. Black with creamy-white markings; 
normally pitted; clothed with very short silvery hairs. 

Head slightly wider than the thorax, black except for the frontal 
eye patches, which are creamy white; clypeal border with a medial 
denticle and two lateral denticles appearing just below and laterad 
of the clypeal process; clypeal process very short and low with a 
tendency to be emarginate on some specimens, subequal in width to 



364 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

the space between the antennae; mandibles with centrally located 
elevations barely evident; antennae normal in form, dark amber 
basally, becoming lighter on the flagellum. 

Thorax black except for two patches on the pronotmn, two small 
patches on the scutellmn, the metanotmn, patches on the tegulae, a 
small patch on the pleuron, all of which are creamy white; enclosure 
deeply and sparsely rugose; mesosternal tubercle small and black; 
legs black except for apical parts of fore- and midfemora aad patches 
on all tibiae, which are creamy white to amber; wings are subhyaline, 
becoming cloudy at the apex. 

Abdomen : tergum 1 with a broad triangular patch ; tergum 2 with 
a broad band cm"ving away from the posterior border of the tergum on 
each side of the center, giving the band a sinuate effect; terga 3 and 
4 with narrower emarginate bands; tergum 5 with an elongate patch; 
venter black, immaculate; pygidium oval but somewhat broader 
basally. 

Male. — Unknown. Possibly it is being confused with the male 
of C. acanthophila Cockerell, C. convergens Viereck and Cockerell, 
or a closely related species. 

Very similar in general appearance to C. acanthophila Cockerell, 
C. convergers Viereck and Cockerell, C. neahminax Scullen, and other 
closely related species. The female of C. butleri Scullen may be 
recognized by the distinct clypeal process, the sinuate band on the 
second abdominal tergum, and the form of the pygidium as well as 
other characters listed in the description. 

Types. — Type female, taken 30 miles south of Safford, Ariz., 
Sept. 24, 195G, on a yellow composite (G. D. Butler), is at the U.S. 
National Museum, no. 66155. Paratypes are as follows: 

Arizona: 9, Canelo, July 30, 1956 (G. D. Butler); ?, 6 mi. E. Douglas, 
Cochise Co., Sept. 8, 1958, at Heplopappus hartwegi (P. D. Hurd) ; 9 , Graham 
Mts., 3500-4500 ft., Aug. 18, 1954 (R. M. Bohart); 9, Portal, July 23-31, 1959 
(K. V. Krombein) ; 9 , Portal, Chiricahua Mts., Aug. 3, 1956 (R. M. Bohart) ; 
3 9 9, 2 mi. NE. Portal, July 30, Aug. 1, 3, 1959 (M. Statham); 9, 4 mi. SE. 
Portal, July 28 1959 (M. Statham) ; 9 , Rosemont, Pima Co., Oct. 9, 1954, at 
Baccharis sp. (F. G. Werner) ; 9 , San Carlos, Sept. 6, 1933 (Parker) ; 9 , Tanque 
Verde, Sept. 12, 1954, at Aplopappus sp. (F. Werner); 9, Tucson, Oct. 19, 1939 
(R. H. Crandall); 9 , Tucson, Nov. 1, 1940 (Bryant). Texas: 9 , Alpine, July 8, 
1952 (H. A. ScuUen). 

Distribution. — Southern Arizona and the one record from Alpine, 
Tex. 

Prey record. — None. 

Plant record. — As Hsted under paratypes. 



WASP GENUS CERCERIS — SCULLEN 365 

7. Cerceris carrizonensis Banks 

Figure 7 
Cerceris carrizonensis Banks, 1915, 403. — Scullen, 1951, p. 1006. 

Max,e. — Length 6 mm. Black with creamy-white markings. 
Tegiilae smooth and not elevated. In color and size it is close to 
acanthrophila Cockerell. Markings of the type are discolored with 
cyanide to some extent. 

This species is known only from the type. It may prove to be a 
synonym, but the writer will retain it as a valid species until more 
positive characters are found to separate these closely related species. 

Type. — The type male, from "Uvalde, Texas, June 18-20, 1930, 
Wickham," is at the Museum of Comparative Zoology, no. 13772. 

8. Cerceris chilopsidis Viereck and Cockerell 

Figures 8, 113a,b,c 
Cerceris chilopsidis Viereck and Cockerell, 1904, p. 136. — Scullen, 1951, p. 1006. 

Female. — Length 10 mm. Color black with creamy-yellow mark- 
ings; normally pitted; clothed w^ith short silvery hairs. 

Head about one-sixth wider than the thorax; black except for large 
frontal eye patches, the clypeus, basal half of the mandibles, and a 
small spot on the front, all of which are creamy yellow, and two 
round spots on the occiput, which are yellow; clypeal border slightly 
extended medially, sinuate; clypeal process scoop shaped but broader 
than long, with the free border black and curved down at the ex- 
tremities; mandibles with two separate, subequal denticles dividing 
the mandible into three subequal parts; antennae normal in form, 
scape largely yellow, peduncle and apical end of scape amber, flagellum 
light below and dark above. 

Thorax black except for two elongate patches on the pronotum, 
two patches on the scutellum, the metanotum, patch below the wing 
base, patches on the mesosternal tubercle, and the tegulae, aU of 
which are yeUow; tegulae smooth and not elevated; enclosiu"e minutely 
laced, with a central groove and pitted at the borders; mesosternal 
tubercles prominent and largely yellow; legs dark amber basally to 
near the apical ends of all femora, remaining parts of legs yellow 
except for patches of amber on all tibiae and the medial tarsi; wings 
subhyaline except for a clouded area at the apex and the stigma, 
which is dark amber. 

Abdomen black except for a semidivided patch on tergum 1 ; sub- 
equal bands on the posterior half of terga 2 to 5 broadly emarginate; 



742-463—65- 



366 



PROCEEDINGS OF THE NATIONAL MUSEUM 



tergum 6 with venter immaculate; pygidium oval with the apical end 
only slightly narrower than the basal, both ends rounded. 

Male. — Unknown. Possibly it is being confused with the male of 
C. convergens Viereck and Cockerell. 

C. chilopsidis Viereck and Cockerell is close to C. convergens Viereck 
and Cockerell, from which it can be separated by the form of the 
clypeal process and the form of the pygidium. The band on tergum 
3 is never broken as it usually is in C. convergens Viereck and Cockerell. 

Types. — The type female of C. chilopsidis Viereck and Cockerell, 
taken at Rincon, N. Mex. (Cockerell), July 5, at flowers of 
Chilopsis saligna, is at the Philadelphia Academy of Natui'al Sciences, 
no. 10375. 




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Figures 7-10. — Localities of: 7, C. carrizonensis Banlcs; 8, C. chilopsidis Viereck and 
Cockerell; 9, C. cleomae Rohwer; 10, C. cockerelli Viereck. 

Distribution. — Southern California, southern Arizona, southern 
New Mexico, southwestern Texas to Laredo, and into Mexico, 

Prey record. — None. 

Plant record. — Alfalfa (Cahfornia), Baccharis glutinosa (Cah- 
fornia), Chiloims saligna (New Mexico), Helianthus annuus (Arizona, 
California), Prosopis sp. (California), Prosopis puhescens (locality not 
recorded), Sasola pestifer (New Mexico), Stromhocarpus puhescens 
(Texas) , Tamarix gallica (California) , Verbesina encelioides (Arizona) , 
Wislisenia sp. (Arizona). 

9. Cerceris cleomae Rohwer 

Figure 9 

Cerceris cleomae Rohwer, 1908, p. 325. — Scullen, 1951, 1006. 

Male. — Length about 5 mm. Black with yellow and yeUowish- 
white markings; punctation and pubescence close to the males of 
the rufinoda-echo group. 



WASP GENUS CERCERIS — SCULLEN 367 

This species is known only from the type, which is in poor condition. 
It closely resembles the males of C. rujinoda Cresson and C. echo 
Mickel, which, so far, are indistinguishable from each other. It may 
prove to be a synonym of one of these species when better characters 
are found for separating these closely related males. The writer is 
retaining it as a valid species for the present. (See discussion under 
C. rujinoda rujinoda Cresson, p. 394.) 

Type. — The type male of C. cleomae Kohwer, from Denver, Colo., 
July 20, 1907, on cleome (Denning), is at the U.S. National Museum, 
no. 25483. 

10. Cerceris cockerelli Viereck 

Figure 10 
Cerceris cockerelli Viereck, 1902, p. 731. — Banks, 1947, p.31. — Scullen, 1951, p. 1006. 

Male. — Length 7.5 mm. Black with creamy-white markings; 
tegulae smooth and not elevated. It is close to C. acanthophila 
Cockerell and C. cleomae Rohwer in color and size. The species is 
known only from the two cotype males. It may prove to be the same 
as some closely related form. Until more positive characters are 
found to separate the males of these closely related species, it will be 
retained as a valid species. 

Type. — The cotype males of C. cockerelli Viereck, from La JoUa, 
Calif., August 1901 (T.D.A. Cockerell), are at the Philadelphia 
Academy of Natural Sciences, no. 10037. 

11. Cerceris conifrons Mickel 

Figures 11, 114a,b,c 

Cerceris conifrons Mickel, 1916, p. 410. — Mickel, 1917b, p. 450. — Banks, 1947, 

p. 31.— Scullen, 1951, p. 1006. 
Cerceris rujinoda crucis H. S. Smith, 1908, p. 370 (in part). — Scullen, 1951, p. 1010. 

Female. — Length 8 to 9 mm. Black with creamy-white and yellow 
markings except anterior parts of abdomen and the propodeum, 
exclusive of the enclosure, which are reddish amber; closely and deeply 
pitted; clothed with very short silvery hairs. 

Head one-fifth wider than the thorax; black except for the frontal 
eye patches, the medial clypeal lobe, patches on the lateral clypeal 
lobes, small patch on the lower frons, two round spots back of the 
compound eyes, the basal half of the mandibles, and the scape, all of 
which are creamy white; clypeal border with three processes, the 
medial one broad and somewhat rounded, the two lateral ones that 
are just laterad of the two curved surface denticles are smaller and 
more pointed; clypeus with a centrally located, low, cone-shaped 
elevation, below which are two black, prominent, curved denticles 
close to the clypeal border; mandibles with two centrally located 



368 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

denticles, the apical one much the larger; antennae normal in form, 
scape yeUow, flagellum light below and dark above. 

Thorax, exclusive of the propodeum, is black except for two elon- 
gate patches on the pronotmn, patches on the pleuron below the 
wing base, and the tegulae, all of which are creamy white; propodeum 
reddish amber except for the enclosure, which is black; tegulae smooth 
and not elevated; enclosure heavily ridged with the ridges subparallel 
to the metanotum and with the lateral ends curving posteriorly, mesal 
groove present; mesosternal tubercle prominent with a trace of yellow 
on the tip; forelegs and midlegs amber to near the apical ends of 
the femora, the remainder of the above legs yellow and amber; hind- 
legs light amber to the apical ends of femora, yeUow changing to 
dark amber beyond. 

Abdomen: terga 1 and 2 reddish amber; tergum 2 with a creamy- 
white band, terga 3 to 5 dark amber with creamy-white bands, tergum 
6 dark amber, immaculate; pygidium broader basally but narrowing 
and truncate at both ends; venter light amber anteriorly, becoming 
dark posteriorly, immaculate. 

Male. — Length 6 to 7 mm. Black with creamy-white and ferru- 
ginous markings; pmictation crowded; pubescence short. 

Head slightly wider than the thorax; black with the exception 
of the face, small evanescent spots back of the eyes, base of mandibles, 
and most of the scape, all of which are creamy white; clypeal border 
with thi-ee indistinct, dentical-like extensions on the medial lobe; 
surface of the medial lobe of the clypeus convex; hair lobes subequal 
in width to the lateral lobes of the clypeus; mandibles without denti- 
cles but with a very low, broad elevation medially; antennae normal 
in form, scape creamy white, fulvous beyond the peduncle, which is 
dark. 

Thorax black except for two widely separated patches on the pro- 
notum, two spots on the scutellum, the metanotum, patch on the 
pleuron, and the tegulae, all of which are creamy white (in some 
specimens the ferruginous of the abdomen spreads more or less onto 
the propodemn) ; tegulae relatively smooth and showing little or no 
elevation; enclosure with heavy horizontal ridges curving laterally 
and a prominent medial groove; mesosternal tubercles absent; fore- 
and midlegs dark fuscous basally to beyond the center of the femora, 
beyond which they are largely creamy white ; hindlegs are dark fuscous 
on the coxae, more or less ferruginous on the trochanters, dark fuscous 
over the femora except the apical end, which is creamy white, the 
tibiae fuscous except for a large creamy-white patch and the tarsi 
ferruginous; wings subhyaline, stigma black. 

Abdomen black except the fust tergum and basal two-thirds of the 
second tergum, which are ferruginous, subequal bands of creamy 



WASP GENUS CERCERIS — SCULLEN 



369 



white on the posterior margins of terga 2 to 6 (in some specimens 
an evanescent, narrow, dark line appears on tergum 2 between the 
yellow band and the ferruginous portion; in some specimens, also, the 
6th tergum is immaculate) ; venter ferruginous over most of sternites 
1 and 2 but dark and immaculate over the remaining sternites; 
pygidium with the sides convex and ends subequal in mdth. 

The males of C. conijrons Mickel may be confused with the males of 
other closely related forms showing ferruginous or reddish coloring 
on the basal segments of the abdomen. The distinctions between 
creamy-white and yellow markings are not always very marked. 

Type. — The type female of C. conifrons Mickel is ai the University 
of Nebraska. 

Distribution. — Except for one female taken at Medicine Hat, 
Alberta, on Aug, 13, 1939, by E. H. Strickland, C. conifrons Mickel 
has been taken only in the western states of Arizona, California, 
Colorado, Nebraska, Nevada, New Mexico, Texas, Wyoming, and 
Utah. It has been taken also in the state of Chihuahua, Mexico. 
(Due to the fact that males of this species sometimes are difficult to 
distinguish from closely related forms, distribution records are based 
only on females.) 

Prey record. — None. 

Plant record. — Aster sp. (Utah), Daucus carota (Utah), Chryso- 
thamnus sp. (New Mexico), Helianthus sp. (Nebraska), Lepidium sp, 
(Arizona), Norta sp. (Utah), Wislizenia sp. (Arizona). 




Figures 11, 12. — Localities of: 11, C. conifrons Mickel; 12, C. convergens Viereck and 
Cockereil. 



12. Cerceris convergens Viereck and Cockereil 

Figures 12, 115a,b,c 

Cerceris convergens Viereck and Cockereil, 1904, p. 136. — Scullen, 1942, p. 188. — 
Banks, 1947, p. 29.— Scullen, 1951, p. 1006. 



370 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ne 

Cerceris rinconis Viereck and Cockerell, 1904, p. 137. 
Cerceris hesperina Banks, 1917, p. 115. 
Cerceris pudorosa Mickel, 1917a, p. 338. 
Cerceris snowi Banks, 1919, p. 84. 

Female. — ^Length 8 to 9 mm. Color black with light yellow 
markings; normally pitted; clothed with short silvery hairs. 

Head one-fifth wider than the thorax; black except for large yellow 
frontal eye patches, the clypeus, and basal part of mandibles, which 
are light yellow; clypeal border slightly extended at the center and 
sinuate; clypeal process low and very short, close to clypeal border, 
divided into two rounded extensions, which are thin at the apex; 
mandibles with two subequal denticles medially located; antennae 
normal in form, scape yellow, peduncle dark amber, flagellum light 
below and dark above. 

Thorax black except for two elongate patches on the pronotum, 
two spots on the scutellum, the metanotum, a patch on each plem'on, 
and the tegulae, all of which are light yellow; tegulae are smooth 
and not elevated; enclosure smooth except for a light medial groove 
and pitted borders; mesosternal tubercle prominent and tipped with 
yellow; legs dark amber basally to near the apical ends of all femora, 
the remainder of each leg being mostly yellow except for an amber 
patch on the apical end of the hindtibiae; wings subhyaline with a 
slight clouded area at the apex. 

Abdomen black except for a semidivided patch on tergum 1, a 
broad band on tergum 2, band on tergum 3, which is usually broken 
into three sections, narrow bands on terga 4 and 5, all of which are 
yellow; venter immaculate; pygidium with the basal end broadly 
rounded and the apical end considerably narrowed to a small rounded 
point. 

Male. — Length 8 mm. Black with yellow to cream markings; 
normally to thinly pitted ; clothed with short silvery hairs. 

Head one-fourth wider than the thorax; black except for two 
yellow frontal eye patches; clypeal border with three centrally lo- 
cated denticles, the medial one slightly more pronounced; mandibles 
without distinct denticles; antennae normal in form, scape yellow, 
peduncle dark amber, flagellum light below and dark above. 

Thorax black except for two elongate patches on the pronotum, 
the metanotum, and the tegulae, which are light yellow; tegulae 
smooth and not elevated; enclosure smooth except for an indistinct 
central groove; mesosternal tubercle absent; legs black basally to 
near the apical ends of all femora, remainder of all legs yellow except 
for black patches on apical ends of hindtibiae and darkened tarsi; 
wings subhyaline except for clouded area at apex. 

Abdomen black except broad band on tergum 2, double broken 



WASP GENUS CERCERIS — SCULLEN 371 

band on tergum 3, and narrow bands on terga 4, 5, and 6; venter 
black, immaculate; pygidium oval with both ends broadly rounded. 

In the male of C. convergens Viereck and Cockerell, the medial 
lobe of the clypeus is usually black but in many specimens more or 
less yellow appears on it. In both sexes it is common to find the 
band on tergum 3 showing slight indentations at the points where 
it is usually broken. In some cases even these indentations disappear. 

C. convergens Viereck and Cockerell is superficially simUar to 
several closely related species. The female can be recognized by 
the clypeal structures and the male by its facial colors and clypeal 
denticles. Both sexes usually show the breaks in the band on the 
third tergum of the abdomen. 

Types. — The type female of C. convergens Viereck and Cockerell, 
from Alamogordo, N. Mex., May 13, 1902 (Viereck), is at the Phila- 
delphia Academy of Natural Sciences, no. 10376. The type female of 
C. hespirina Banks, taken at Yakima, Wash., July 1882 (Samuel 
Henshaw), is at the Museum of Comparative Zoology, Harvard 
University, no. 10,031. No allotype male was indicated. The type 
female and allotype male of C. pudorosa Mickel, both taken at Auburn, 
Calif., Sept. 20, 1916 (L. Bruner), are at Nebraska State University, 
The type female of C. rinconis Viereck and Cockerell, from Rincon, 
N. Mex., July 5, at flowers of Chilopses saligna, is at the Philadelphia 
Academy of Natural Sciences, no. 10386. The lectotype male of C. 
snowi Banks, from Tucson, Ariz. (Snow), is at the Museum of Com- 
parative Zoology, Harvard University, no. 13764. 

Distribution. — This is one of the most widely distributed species 
of the Pacific Slope. It ranges east through the Southern States to 
eastern Texas. Records of males taken at Sherman Co., Kans., 
and Yankton, S. Dak., may be open to question. 

Prey record. — Bruchus sp., Alhambra, Calif,, July 14, 1921 (A. O. 
Larson) , 

Plant record. — Acacia sp. (Arizona, New Mexico), Acacia 
angustissima (Arizona), Acacia greggii (Ai'izona), Achilles sp. (Idaho), 
Adenostom sp. (California), alfalfa (Arizona, Utah), Aplopap2^us sp. 
(Arizona), Asclepias sp. (Arizona, California), Atriplex sp. (Utah), 
Baccharis sp. (Arizona), Baccharis douglasii (California), Baccharis 
emeryi (California), Chilopsis saligna (New Mexico), Chilopsis linearis 
(Arizona), Chrysothamnus sp. (California, Utah), Cissus trijoliata 
(Arizona), Cleomelia sp. (Idaho), Cleome lutea (Oregon), Condalia sp. 
(Arizona), cotton (Arizona), Croton californicus (California). 
Eriogonum sp. (Idaho, Oregon, Utah), Eriogonum (annual, California), 
Eriogonum eletum (Oregon), Eriogonum fasciculatum (California), 
Eriogonum fasciculatum 'polifolium (California), FJriogonum gracile 
(California), Eriogonum injlatum (California), Gaillardia amhlyodon 



372 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

(Texas), Gnaphalium sp. (California), Gnaphalium beneolens (Cali- 
fornia), Gutierrezia sarothrae (California), Helianthus sp. (Idaho), 
Helianthus annuus (California), Larrea sp. (California), Polygonum 
sp. (Arizona), Polygonum lapathvfolium (California), Salix taxifolia 
(Arizona), Salsola kali (Utah), Sapindus saponari (Arizona), Solan 
elegan (Aiizona), Solidago sp. (California, Oregon), Tamerix gallica 
(California), Xanthium sjnnosum (California). 

13. Cerceris crandalli, new species 

Figures 13, 116a,b,c 

Female. — ^Length 8 mm. Black to dark amber with creamy-white 
and light yellow markings; normally pitted and clothed with very 
short silvery hairs. 

Head about one-fifth wider than the thorax; black except for large 
frontal eye patches, small spot on the clypeal process, oval spots back 
of the compound eyes, elongate patches on the vertex back of the 
ocelli, patches on the mandibles and the scape, all of which are creamy 
white; clypeal border with a broad rounded slight extension mesad, 
on each side of which is a small denticle ; clypeal process scoop shaped 
but much broader than long with the free border deeply emarginate; 
mandibles with two distinct, centrally located denticles, the apical 
one much the larger and more rounded; antennae normal in form, 
scape creamy white, peduncle and flagellum amber. 

Thorax black except for two elongate patches on the pronotum, 
two spots on the scutellum, the metanotum, patches on the pleuron 
below the wing base, the tegulae, and the apex of the mesosternal 
tubercle, all of which are creamy white; tegulae smooth and not 
elevated; enclosure deeply ridged subparallel to the metanotum; 
mesosternal tubercle medimn in size; legs dark amber to near the 
apical ends of the femora, the remainder of the leg parts are yellow 
to light amber except for dark patches on all tibiae; wings subhy aline, 
cloudy at the apex. 

Abdomen black except for a small patch on tergum 1, a broad band 
on the posterior half of tergum 2, narrow bands on terga 3 and 4, 
and a small patch on tergum 5, aU of which are light yellow; venter 
black, immaculate; pygidium with sides convex, basal end about one- 
half the greatest widtli, and the apical end broadly rounded. 

Male. — Length 7 mm. Black with creamy-white to light yellow 
markings; punctation average; pubescence short. 

Head about one-fiftti wider than the thorax; black except for large 
frontal eye patches and the scape, which are light yellow; clypeal 



WASP GENUS CERCERIS — SCULLEN 373 

border with three small denticles, the medial one the largest; hair 
lobes subequal to the lateral clypeal lobes; mandibles without denticles; 
antennae normal in form, fulvous but slightly darker above. 

Thorax black except for a divided band on the pronotum, the 
metanotum, a minute spot on the pleuron below the wing, and a patch 
on the tegulae, all of which are creamy white; tegulae smooth and not 
elevated; enclosure deeply ridged subparallel to the metanotum; meso- 
sternal tubercles absent; legs black basally to near the apical ends of 
the femora, which are creamy white; tibiae and other apical segments 
creamy white except for the apical half of the 3rd tibia, which is dark 
I'uscous to black; the light tip of the 3rd femur cut off from the dark 
area abruptly at right angles to the femur as in C. acanthojphila 
Cockerell; wings subhyaline but clouded apically; stigma dark. 

Abdomen black with a small semidivided spot on tergum 1, a broad 
band on the posterior half of tergum 2, narrow bands on the posterior 
margins of terga 3, 4, and 5, and a small patch on tergum 6, all of 
which are creamy white; venter black, immaculate; pygidium with 
sides convex and both ends rounded. 

The male is very near that of C. acanthophila Cockerell, from which 
it may be distinguished by a creamy-white face and the relatively 
smooth enclosure of the latter species. 

Types. — The type female from Tucson, Ariz., June 13, 1938, 
(R. H. Crandall), is deposited at the U.S. National Museum, 
no. 66156. The allotype male, from 8 miles southeast of Elota, 
Sinaloa, Mexico, 1962 (F. D. Parker), is deposited at the University 
of Cahfornia, Davis, Calif. 

Arizona: ? , 10 mi. E. Gila Bend, June 15, 1955 (G. D. Butler); 4 ? 9 , 2 mi. 
NE. Portal, Cochise Co., July 30- Aug. 5, 1959, Baccharis (M. Statham); $, 
Sahuarita, July 5, 1956 (G. D. Butler) ; 3 9 9 , Toltec, Pinal Co., June 14-15, 
1953 (T. R. Haig); 9, Tombstone, July 27, 1955, Mortonia scalrella (Butler- 
Werner); 2 9 9, Tucson, June 10, 13, 1938 (R. H. Crandall); 9, Tucson (F. H. 
Snow). Texas: 9, Boquillas, Brewster Co., June 30, 1928 (F. M. Gage); 9, 
20 mi. N. El Paso, June 19, 1942 (H. A. Scullen). Mexico: Sinaloa: 59 9, 
16c? d^, 8 mi. SE. Elota, May 18-19, 1962 (F. D. Parker, L. A. Stauge); 2 9 9, 
3cf d^, Cocorit, May 23, 1962 (F. D. Parker, L. A. Stange) ; 9 , 2cf cf', 5 mi. S. 
Magdalena, May 25, 1962 (F. D. Parker, L. A. Stange); 9 , d^, 5 mi. W. Santa 
Ana, June 10, 1962 (F. D. Parker). 

Distribution. — Southern Arizona, western Texas, and northwest- 
ern Mexico. 

Prey record. — None. 

Plant record. — Baccharis (Arizona), Mortonia scalrella (Arizona). 



374 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 13-15. — Localities of: 13, C. crandalli Scullen; 14, C. crotonella Vlereck and Cock- 
erell; 15, C. echo echo Mickel. 



14. Cerceris crotonella Viereck and Cockerell 

Figures 14, 117a,b,c 
Cerceris crotonella Viereck and Cockerell, 1904, p. 139. — Scullen, 1951, p. 1007. 

Female. — Length 8 mm. Color black and reddish amber, marked 
with creamy white; normally pitted; clothed with very short silvery 
hairs. 

Head one-fourth wider than the thorax; black except for large 
frontal eye patches, the clypeus, basal half of mandibles, patch on 
the front, round spots back of the compound eyes, and the scape, all 
of which are creamy white; clypeal border with five subequally spaced 
low denticle-like processes; process on the clypeal surface extended 
to form a truncate cone with the end slightly broadened and black; 
mandibles with two separate, subequal denticles, medially located; 
antennae normal in form, scape creamy white, peduncle and flagellum 
light amber below, darker above. 

Thorax, exclusive of the propodeum, black except for two elongate 
patches on the pronotum, two patches on the scutellum, the meta- 
notum, patches on the pleuron below the wing attachments, and the 
tegulae, all of which are creamy white; tegulae smooth and not 
elevated; propodeum reddish amber except the enclosure, which is 
black ; enclosure deeply pitted along the border and along the central 
groove mth a few incomplete ridges next to the mentanotum; meso- 
sternal tubercle absent; legs dark amber basally to a little beyond 
the middle of the femora; apical ends of femora, the fore- and mid- 
tibiae, and the hindtibiae in part, creamy white; tarsi amber, becoming 
darker on the mid- and hindlegs; mngs subhyaline. 

Abdomen: terga 1 to 3 reddish amber with narrow creamy-white 
bands; terga 4 to 5 black with creamy- white bands; tergum 6 black, 



WASP GENUS CERCERIS — SCULLEN 375 

immaculate; venter reddish amber anteriorly, becoming dark amber 
posteriorly; pygidium broader basally and rounded, the apical end 
narrowing and truncate. 

Male. — Length 6 to 7 mm. Black with creamy- white and fer- 
ruginous markings; punctation more widely spaced than average; 
pubescence short. 

Head slightly wider than the thorax; black except for the face, 
base of mandibles, and scape, all of which are creamy white; clypeal 
border with three subequal low denticles on the medial lobe; hair 
lobes extend over about three-fourths of the lateral clypeal lobes; 
surface of the medial lobe convex; mandibles without denticles; 
antennae normal in form, scape largely creamy white, flagellum 
largely fulvous. 

Thorax black except for two elongate patches on the pronotum, 
two large spots on the scuteUum, the metanotum, a small patch on 
the pleuron, and the tegulae, all of which are creamy white; tegulae 
low and smooth; enclosure with a pitted medial groove and deeply 
rugose laterally; mesosternal tubercle absent; legs black to the apical 
ends of the femora, beyond which they are creamy white except for 
the apical segments of the tarsi, which become darker; the hindtibiae 
and the hindtarsi dark except for a large, creamy patch on the tibiae ; 
wings are subhyaline, stigma black. 

Abdomen black or dark fuscous except the first segment and the 
basal part of the second segment, which are ferruginous, and the 
following creamy-white markings: broad bands on terga 2 and 3, 
which have the anteriolateral corners darkened, narrow bands on 
terga 4 and 5, an evanescent patch on tergum 6; venter with minute, 
creamy spots laterally on sternites 3 and 4; pygidium with sides 
convex and the apical end slightly smaller than the basal end. 

The male of crotonella Viereck and Cockerell is close to the males 
of C. conifrons Mickel and C. hridwelli ScuUen. They are distinguished 
from conifrons by the more closely crowded punctation and narrow 
light bands on the abdomen of the latter. They are distinguished 
from hridwelli by having only two broad bands on the abdomen 
while the latter has three. The surface punctation is sunilar to 
hridwelli. 

Type. — The type female of C. crotonella Viereck and Cockerell, 
taken at Las Cruces, N. Mex., on Sept. 25, 1895, Croton neomexicanum 
(Cockerell), is at the Philadelphia Academy of Natural Sciences, 
no. 10039. 

Distribution. — Southern Arizona, New Mexico, and southwestern 
Texas. 

Prey record. — None. 



376 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lis 

Plant record. — Acacia augustissima (Arizona), Chilopsis sp. (New- 
Mexico), Crotonella neomexicanum (New Mexico), Larrea tridentata 
(Texas), Opuncta sp. (Texas), Pedis pepposa (New Mexico). 

15a. Cerceris echo echo Mickel 

Figures 15, 118a,b,c 

Cerceris echo Mickel, 1916, p. 412.— Mickel, 1917b, p. 453.— ScuUen, 1951, p. 1007. 
Cerceris rufinoda crucis H. S. Smith, 1908, p. 370 (in part). 

Female. — Length 9 to 10 mm. Color black and reddish amber 
with yellow markings; closely and deeply punctate; clothed with 
very short sUvery hairs. 

Head one-sixth wider than the thorax, black except large lateral 
eye patches, small spots back of the compound eyes, basal end of 
mandibles, and patch on scape, which are yellow; clypeal border 
with two widely separated broad extensions, between which is a 
single denticle, and laterad of each extension is a low denticle-like 
process; clypeal surface process is in the form of a low transverse 
ridge wdth the extremities dentiform; mandibles with one centrally 
located denticle, more apicad of which is a low rounded elevation; 
antennae normal in form, scape yellow to light amber, peduncle 
amber, flagellum light amber below, dark above. 

Thorax, exclusive of the propodeum, black except for two elongate 
patches on the pronotum, two spots on the metanotum, and patch 
on the tegulae, all of which are yellow; tegulae sparsely pitted and 
slightly elevated; propodeimi dorsally reddish amber except for the 
enclosure, which is black; enclosure deeply pitted and rugose with a 
prominent central groove; mesosternal tubercle small and black; legs 
largely amber basally, becoming more yellow beyond the middle of 
the femora; wings slightly clouded. 

Abdomen: tergum 1 reddish amber; terga 2 to 5 black with bands 
of light yellow, that on tergum 2 broader and not emarginate as are 
the others; venter black except for the first stemite, which is reddish 
amber; pygidium oval, narrowing, and rounded at both ends. 

Male. — Unknown. It may be confused with closely related 
species. (See note under rufinoda rufinoda Cresson, p. 394.) 

Type. — The type female, taken at Monroe Canyon, Sioux Co., 
Nebr., Aug. 4, 1908 (C. H. Gable), is at the University of Nebraska. 

Distribution. — Widely distributed but not collected in any great 
numbers over the Central Plains States from southern Canada to the 
Mexican border. Recorded from Alberta, Idaho, Montana, North 
Dakota, South Dakota, Minnesota, Iowa, Nebraska, Kansas, 



WASP GENUS CERCERIS — SCULLEN 377 

Colorado, California, Utah, Arizona, New Mexico, and Texas. An 
isolated record from Independence Lake, Sierra Co., Calif., July 20, 
1954 (R. C. Blaylock), is of special interest. 

Prey record. — None. 

Plant record. — Aster sp. (Kansas), Baccharis sp. (Texas), Chryso- 
thamnus sp. (Utah), Eupatorium serotinun (Texas), parsnip (Texas), 
Pastinaca sativa (Texas), Solidago sp. (New Mexico), Thelesperma 
gracile (Kansas). 

15b. Cerceris echo atrata, new subspecies 

Figure 16 

Female. — This is the same in structm'e and color pattern as C. 
echo echo Mickel except that the markings are light yellow and it 
lacks the reddish amber on the propodeum and the first abdominal 
segment so characteristic of the latter. 

Male. — Unknown. It may be confused with the male of C. 
rufinoda rujlnoda Cresson, with which it is closely related. 

This is the eastern form of C. echo Mickel. From manuscript 
labels by Banks found on some specimens, it is evident he had at 
one time expected to describe this form as a new species under the 
name of nigroris. 

Types. — The type female, from Camden Co., N.J., July 12, 1891 
(Wm. J. Fox), is at the Philadelphia Academy of Natural Sciences, 
no. 5039. Paratypes are as follows: 

Georgia: 4? ?, Atlanta, June 14, 1936 (P. W. Fattig) ; ?, Cedartown, June 
7, 1936 (P. W. Fattig). Illinois: 9, Carlinville (Charles Robertson, 6129); 9, 
Meredosia, Aug. 22, 1917, sand pit. Iowa: ?, Sioux City, Aug. 11, 1928 (C. N. 
Ainslie). Kansas: ? , Baldwin, July (Bridwell) ; ? , Douglas Co., 900 ft., (F. H. 
Snow); 29 9, Riley Co., July 14, 25 (Pepenoe, 5342). Massachusetts: 9, 
Forest Hills, July 1913 (F. X. Williams); 9, Hopkinton Forest, Aug. 22; 9, 
Reading, July 14, 1935, Umbelliferae flowers (Richard Dow). Maine: 9, 
Desert of Maine, Aug. 13, 1950. North Carolina: cf , (Cresson). New Jersey, 
69 9, Camden Co., July 12, 1891 (Wm. J. Fox). New York: 9, Ardsley, 
Westchester Co., Aug. 9, 1958 (G. R. Ferguson) ; 9 , Bedford, Westchester Co., 
July 20, 1957 (G. R. Ferguson); 29 9, Hartsdale, Westchester Co., July 14, 
1957 (G. R. Ferguson); 29 9, Pound Ridge, Westchester Co., July 21, Aug. 17, 
1957 (G. R. Ferguson). Virginia: 9 , East Falls Church, July 20 (S. A. Rohwer) ; 
9, Vienna, Aug. 11, 1935 (J. C. Bridwell). Wisconsin: 9, Genoa, Vernon Co., 
July 7-12, 1911 (53145). 

Distribution. — Limited through the Eastern States east of the 
Mississippi River. 

Prey record. — None. 

Plant record. — Umbelliferae (Massachusetts). 



378 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 16, 17. — Localities of: 16, C. echo atrata Scullen; 17, C. finitima finitima Cresson. 



16a. Cerceris finitima finitima Cresson 

Figures 17, 119a,b,c 
Cerceris finitima Cresson, 1865, p. 125. — Packard, 1866, p. 61. — Cresson, 1887, 

p. 282.— Schletterer, 1887, p. 491.— Robertson, 1890, p. 200; 1891, p. 570; 

1892b, p. 108; 1892c, p. 274; 1894a, pp. 455, 457.— Dalla Torre, 1897, p. 460.— 

Bridwell, 1898, p. 209.— Ashmead, 1899, p. 295.— Viereck and Cockerell, 1904, 

p. 138.— Smith, H. S., 1908, p. 370.— Banks, 1912a, p. 27.— Stevens, 1917, p. 

422.— Rohwer, 1917, p. 244.— Mickel, 1917b, p. 453.— Rau, 1928, p. 338.— 

Strandtmann, 1945, p. 312.— Scullen, 1951, p. 1007. 
Cerceris finitima var. nigroris Banks, 1912a, p. 27. — Mickel, 1917b, p. 453. — 

Scullen, 1951, p. 1008. 

Female. — Length 7 to 8 mm. Color black and yellow, closely 
and deeply pitted, clothed with very short silvery hairs. 

Head one-fourth wider than the thorax, black except large frontal 
eye patches on the face, the clypeus, patch on front above the clypeus, 
basal part of the mandibles, the scape, spots back of the compound 
eyes, all of which are yellow; markings on face lighter than other 
markings ; clypeal border with two widely separated denticles ; clypeal 
surface with a very low, broad ridge ending laterally in two slightly 
extended subhy aline points appearing just above and mesad of the 
two dark border denticles; mandibles with two subequal denticles 
distinctly separated and medially located; antennae normal in form, 
scape yellow, peduncle dark amber, flagellum light amber below and 
darker above. 

Thorax black except for two elongate patches on the pronotum, 
two patches on the scutellum, the metanotum, a dot on the pleuron 
below the wing base, and the tegulae, all of which are yellow; tegulae 
elevated and pitted; enclosure transversely rugose; mesosternal 
tubercles small and black ; legs dark amber to middle of the femora or 
beyond, yellow beyond to the tarsi, which are light amber; wings 
subhyaUne except the stigma, which is dark amber. 



WASP GENUS CERCERIS — SCULLEN 379 

Abdomen with deeply emarginate bands of yellow on terga 1 to 5, 
wider on tergum 3; venter immaculate; pj^gidium oval with both 
ends rounded, the basal end more acute. 

Male. — Length 6 to 7 mm. Color black and yellow, closely and 
deeply pitted, clothed with very short silvery hairs. 

Head about one-fifth wider than the thorax; black except for the 
face, which is yellow; clypeal border with three subequal, centrally 
located denticles; mandibles smooth; antennae normal in form, scape 
yellow, peduncle dark amber and fiagellum light amber below, dark 
above. 

Thorax black except for elongate patches on the pronotum, two 
patches on the scutellum, the metanotum, small spots on the pleuron 
below the wing bases and the tegulae, aU of which are yellow; tegulae 
elevated and pitted; enclosure deeply rugose transversely; mesopleural 
tubercle absent ; legs dark amber basally to the middle of the femora 
or a little beyond, yellow beyond, becoming light amber on the 
tarsi; wings subhy aline. 

Abdomen black except for a patch on the first tergum, broad band 
on tergum 2, narrow bands on terga 3 to 6 emarginate, which are 
yellow; venter dark amber with a tendency for yellow spots to appear 
laterally on sternites 3 to 5; pygidium oval Avith the basal end rounded, 
apical end more truncate. 

Types. — The type female of C.finitima Cresson, from Illinois (Dr. 
Lewis), is at the Pliiladelphia Academy of Natural Sciences, no. 1948. 
The type male of C.Jinitima var. nigroris Banks, from Falls Church, 
Va., is at the Museum of Comparative Zoology, Harvard University, 
no. 13768. 

The writer does not consider the variety nigroris Banks distinctive 
enough to recognize it as a subspecies. 

Distribution. — Widely distributed throughout the states east of 
the Kocky Mountains except in New England. It ranges from 
southern Canada to the Mexican border. It is found also in Cali- 
fornia, from Davis south, and into Arizona and New Mexico. 

Prey record. — Chaetocnema pulicaria, a flea beetle (Chrysomeli- 
dae), Columbus, Ohio (Strandtmann, 1945, p. 312). 

Plant record. — Acacia sp. (Texas), Asclepias sp. (Arizona, 
Nebraska, New Mexico), Aster sp. (Kansas), Bijora americana (Texas), 
Cassia sp. (Texas), celery (Utah), Cleome serrulata (Nebraska), clover 
(Illinois), cowpeas (Arkansas), Croton calij'ornicus (California), Daucus 
carota (Ohio), Dichro phyllum (Colorado), Erigeron quercijolius 
(Florida), Grindelia sp. (South Dakota), Orindelia squarrosa (Utah), 
Helianthus pumilus (Colorsido) , Kuhnistera oligophylla (North Dakota), 
Melilotus alba (Nebraska), Monarda citriodora (Texas), pea vine 
(Ohio), Polygonum auberti (California), Sambucus canadensis (Ohio), 



380 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Solidago sp. (Colorado, Iowa, Kansas, Nebraska, Texas), Telesperma 
gracile (Utah). 

16b. Cerceris finitima citrina, new subspecies 

Figure 18 

Female. — Length 6 to 7 mm. Color black with yellow markings 
covering over fifty per cent of the body surface; deeply and closely 
pitted ; clothed with short silvery hairs. 

Head about one-fifth wider than the thorax; black but with the 
yellow marldngs covering most of the face, basal half of the mandibles, 
scape, and large irregular patch extending from one gena through 
vertex to the opposite gena ; clypeal border with two widely separated 
denticles ; clypeal surface with a very low, broad ridge ending laterally 
in two slightly extended subhyaline points appearing just above and 
mesad of the two dark border denticles ; mandibles with two centrally 
located but distinctly separated denticles, the more apical one much 
the larger and pointing distally; antennae normal in form, the scape 
yellow, peduncle amber and the flagellum light amber below but 
darker above. 

Thorax black except for a broad band on the pronotum, two short 
stripes on the scutum, the scutellum, the metanotum, large patches 
on the propodeum, two large patches on each pleuron, two small 
spots on the sternum, and the tegulae, all of which are yellow; tegulae 
elevated and lightly pitted; enclosure black, transversely rugose; 
mesostemal tubercle present, partly yellow; legs largely yellow except 
for small dark patches on the basal areas of each leg ; wings subhyaline 
except the stigma, which is amber. 

Abdomen with broad yellow bands on terga 1 to 5, those on terga 
3 to 5 somewhat emarginate; pygidium oval, broader basally, apical 
end rounded, basal end truncate; venter dark amber except for a 
mesal and two lateral yellow patches on stemites 2 to 4. 

Male. — Length 6 to 7 mm. Color black with yellow marldngs; 
deeply and closely pitted ; clothed with short silvery hairs. 

Head about one-third wider than the thorax; black except entire 
face, elongate patches on the genae, the vertex, basal two-thirds of 
the mandibles, and the scape, all of which are yellow; border of clypeus 
sinuate with two lateral and one medial slight extensions; clypeal 
surface convex; mandibles smooth; antennae normal in form, scape 
yellow, peduncle amber, flagellum light amber below, darker above. 

Thorax black except for two elongate patches on the pronotum, 
the scutellum, the metanotum, large triangular patches on the pro- 
podeum, two patches on the pleuron, and the tegulae, all of which 
are yellow; mesostemal tubercle absent; tegulae elevated and lightly 



WASP GENUS CERCERIS — SCULLEN 



381 



pitted; enclosure heavily rugose transversely; legs yellow except for 
small patches of amber on the following: coxae, all femora, the 
hindtibiae, and hindtarsi ; wings subhyaline except the stigma, which 
is dark amber. 

Abdomen vtdth broad yellow bands on terga 1 to 6, those on terga 
5 and 6 emarginate; venter amber with yellow bands on sternites 2 
to 4 and two yellow patches on stemite 5 ; pygidium with sides convex 
and both ends subtruncate. 

The extent of the yellow markings vary in both sexes. This is 
especially true of the yellow line on the genae, venter, and scutellum. 
Smaller patches may disappear. The subspecies citrina has far more 
yellow than the typical form C . finitima finitima Cresson. 

Types. — The type female, from Riverside, Cahf., Aug. 29, 1926, on 
Polygonum lapathifolium (P. H. Timberlake), and the allotype male, 




Figures 18-20. — Localities of: 18, C. finitima citrina Scullen; 19, C finitima vierecki Banks; 
20, C. irene Banks. 



from Riverside, Calif., Aug. 26, 1927 (P. H. Timberlake), are at the 
Cahfornia Academy of Sciences. Paratypes are as follows: 

Califoenia: 9, Antioch, Sept. 18, 1938 (J. W. MacSwain); ?, Claremont 
(Baker); 9, Colton, Aug. 10, 1951 (J. C. HaU) ; 2 99, Davis, Aug. 24, 1939 
(G. E. Bohart) ; 9 , Davis, July 11, 1951 (E. I. Schlinger) ; cf, Davis, July 11, 1954 
(J. 0. Downey); cf, Davis, Aug. 20, 1950 (R. C. Bechtel); cT, Lindsay, 1909, on 
Asclepias (W. A. Davidson) ; 9 , Patterson, Stanislaus Co., June (T. F. Leigh) ; 
9 , Riverside, Sept. 23, 1924, on annual Eriogonum (P. H. Timberlake) ; 9 , 
Riverside, Aug. 16, 1927, on Eriogonum gracile (P. H. Timberlake); 9 , Riverside, 
Aug. 29, 1926, on Polygonum lapathifolium (P. H. Timberlake) ; 9 , Riverside, 
Oct. 1, 1928, on Euphorbia albomarginata (P. H. Timberlake) ; 9 , Riverside, 
July 17, 1940 (C. Daramers) ; cf , Riverside, Sept. 20, 1933, on Erigeron linifolius 
(P. H. Timberlake); d', Santa Barbara, June 1907 (W. M. Gififard); cf, Visalia, 
1927, on privet (P. R. Jones). 
742-463—65 — —4 



382 PROCEEDINGS OF THE NATIONAL MUSEUJVl vol. us 

Distribution. — Recorded only from Davis and southern California. 

Prey records. — None. 

Plant records. — As indicated under type material. 

16c. Cerceris finitima vierecki Banks, new status 

Figure 19 

Cerceris vierecki Banks, 1947, p. 30. — ScuUen, 1951, p. 1011. 
Cerceris finitima vierecki Scullen, 1960, p. 80. 

Female. — Length 8 mm. Colors black with creamy-white mark- 
ings; finely and moderately pitted; pubescence average. 

Head about one-fifth wider than the thorax; black except for face 
and round spots back of the compound eyes, which are creamy white; 
clypeal border with two widely separated pointed extensions; clypeal 
surface slightly convex without special elevations; mandibles with 
two subequal denticles distinctly separated, the apical one much the 
larger; antennae normal in form, scape creamy white, peduncle dark 
amber, flagellum light amber below and darker above. 

Thorax black except for two elongate patches on the pronotum, 
two patches on the scutellum, the metanotum, large patch on the 
pleuron below the wing attachment, and patches on the tegulae, all 
of which are creamy white; tegulae more smooth and less elevated 
than in C. finitima finitima Cresson ; enclosure smooth except for the 
central groove, which is pitted, and the lateral borders, which are 
slightly rugose; mesosternal tubercle extremely inconspicuous; legs 
black basally to beyond the middle of the femora, creamy white 
beyond except for an amber patch on the mesal side and apical end 
of the hindtibiae and the tarsi, which are darker; wings subhyaline 
except the stigma, which is dark amber. 

Abdomen black except for a patch on tergum 1, and emarginate 
bands on terga 2 to 5, which are creamy white; venter black, immac- 
ulate; pygidium oval with both ends rounded. 

Male. — The males taken in the same localities as the females 
of C. finitima vierecki are of the typical form and color pattern of C. 
finitima finitima but the markings tend to be more creamy white as 
is true of the females. 

Type. — The type female of Cerceris vierecki Banks, taken at Tempe, 
Ariz., Aug. 1 (Bequaert), is at the Museum of Comparative Zoology, 
no. 23544. 

Distribution. — Largely throughout the Southwestern desert area 
of Arizona, New Mexico, and Texas; rare in Colorado and Utah. 

Prey record. — None. 

Plant record. — Acacia sp. (Arizona), Asclepias subverticillata 
(Arizona), Baccharis glutinosa (Arizona), cotton (New Mexico), 



WASP GENUS CERCERIS — SCULLEN 383 

Eriogonum abertianum neomexicana (Arizona), yellow composit 
(Arizona) , 

17. Cerceris irene Banks 

Figures 20, 120a,b,c 
Cerceris irene Banks, 1912a, p. 26— ScuUen, 1951, p. 1008. 

Female. — ^Length 7 to 8 mm. Color dark amber, reddish amber 
and yellow; closely and deeply pitted; clothed with very short silvery 
hairs. 

Head one-fifth wider than thorax; amber except face, basal half 
of mandibles, scape, patch on gena, and elongate patches on vertex, 
which are yellow; ocellar area dark amber; clypeal border with one 
central denticle-like process, to each side of which is a broad extension 
of the clypeal border; clypeal process with a broad rounded base 
narrowing to a truncate point; mandibles with two centrally located 
denticles, the apical one the larger; antennae normal in form, scape 
yellow, peduncle and flagellum light amber. 

Thorax reddish amber below to dark amber or black above except 
two elongate patches on the pronotum, two patches on the scutellum, 
the metanotum, patch on the pleuron, and the tegulae, all of which 
are yellow somewhat infused with light amber; tegulae elevated and 
somewhat pitted; enclosure reddish amber to dark amber, deeply 
rugose transversely; propodeum reddish amber to amber; mesosternal 
tubercle present; legs reddish amber; wings subhyaline except the 
stigma, which is darker. 

Abdomen, tergum 1 reddish amber with a broad emarguiate yellow 
to amber patch; terga 2 to 5 with broad emarginate bands of yellow 
infused with amber; venter reddish amber to dark amber with reddish 
amber bands on sternites 2 to 3; pygidium with convex sides and 
truncate ends. 

Male. — The male of irene has not been isolated. It is probably 
being confused with the males of closely related species (see note under 
rufinoda rufinoda Cresson, p. 394) . 

C. irene Banks is extremely variable in color shade and the extent 
of color markings. The ground color varies from black to medium 
amber. The yellow patches on the pronotum and the scutellum may 
fuse to form bands on the respective parts. 

Type. — The type female, from Fedor, Lee Co., Tex., June 25 
(Birkman), is at the Museum of Comparative Zoology, Harvard 
University, no. 13781. 

Distribution. — Except for the one Minnesota record, C. irene 
Banks is found only in the Western and South Central States of 
Wyoming, Colorado, Kansas, Oklahoma, Texas, and Louisiana. 
Specimens are as follows: 



384 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Colorado; 9 , Larimer Co., Aug. 13, 1952 (R. R. Dreisbach). Kansas: 29 9, 
Blackjack Creek, Pottawatomie Co., July 2, 14, 1953 (Evans, Lin, Yoshimoto) ; 
9 , Blackjack Creek, July 9, 1954 (H. E. and M. A. Evans); 9 , Madero, June 21, 
1936 (W. O. Field); 9, Madero, July 7, 1953 (H. E. Evans); 9, Riley Co., Sept. 
12 (J. B. Norton) ; 3 9 9 , Riley Co., Sept. 17, 19 (Popenoe) ; 9 , Reno Co., Aug. 
13-20, 1917; 9, Stafford Co., Salt Flat Area, July 11, 1957 (H. E. and M. A. 
Evans); 9, Wallace Co., 3440 ft. (F. X. Williams). Louisiana: 9, Keatchie, 
June 14, 1905. Minnesota: 9, Fridley Sand Dunes, Anoka Co., July 22, 1927 
(R. W. Dawson). Oklahoma: 9, Alfalfa Co., Aug. 10, 1932 (Deonier and 
Pritchard). Texas: 9, Ballinger, Aug. 3, 1931 (R. H. Painter); 9, Bexar 
Co., May 1, 1929 (H. B. Parks); 9, Brooks Co., July 25, 1928 (R. H. Beamer) ; 
9, Brooks Co., July 25, 1928 (J. G. Shaw); 9, Corpus Christi, June 28, 1942 
(E. S. Ross); 9, Fedor, Lee Co., May 7, 1909; 9, Fedor, July 23, 1897 (Birk- 
mann); 9, Fort Worth, May 31, 1937 (Heard and Berjim); 49 9, Gillette, 
Karles Co., July 25, 1917; 9 , KendaU Co., July 22, 1928 (R. H. Beamer) ; 4 9 9 , 
Lee Co., May 30, Sept. 21, 1906; 9, McBade, Bastrop Co., May 12, 1934 (J. E. 
Gillaspy); 9, Neuecest, April 28, 1896 (Morlatt); 49 9, Port Isabel, Cameron 
Co., June 23-27, 1956 (H. E. Evans and E. G. Matthews) ; 9 , Reno Co., Aug. 
13-20, 1917; 9, Rosser, July 6, 1905 (C. R. Jones); 9, Victoria, June 24, 1917; 
9, Wallace Co., 3440 ft. (F. X. Williams). Wyoming: 9, Summit, 8835 ft., 
Aug. 16, 1940 (H. E. MiUiron). 

Prey record. — None. 

Plant record. — Gaillardia amhlyodon (Texas). 

18a. Cerceris kennicottii kennicottii Gresson 

Figures 21, 121a,b,c 

Cerceris kennicottii Cresson, 1865, p. 128. — Packard, 1866, p. 63. — Cresson, 
1872, p. 231; 1887, p. 282.— Schletterer, 1887, p. 495.— Robertson, 1890, p. 
200; 1894a, pp. 457, 471; 1896b, p. 73.— Dalla Torre, 1897, p. 464.— Bridwell, 
1898, p. 209.— Ashmead, 1899, p. 295.— Banks, 1912a, p. 25.— Mickel, 1917b, 
p. 452.— Hendrickson, 1930, p. 159.— Banks, 1947, p. 30.— Scullen, 1951, 
p. 1008. 

Cerceris eriogoni Viereck and Cockerell, 1904, p. 139. — Scullen, 1951, p. 1007 
(new synonymy). 

Female. — Length 8 to 10 mm. Black with yellow markings; 
normally pitted; clothed with very short silvery hairs. 

Head about one-fifth wider than thorax; black except for large 
frontal eye patches, patch on clypeal process, basal part of mandibles, 
and scape, all of which are yellow; clypeal border with five equally 
spaced small processes, the central one denticle-like, the two lateral 
ones also denticle-like but smaller, the other two somewhat broader; 
clypeal process broad at the base, converging distally to a semitruncate 
apical end, which is sinuate, showing a small medial and two small 
lateral extensions; mandibles with two subequal, centrally located 
denticles; antennae normal in form, scape light yellow in front, 
peduncle dark amber, flagellum light amber below and dark above. 

Thorax black except for two elongate patches on the pronotum, 
band on the scutellum, minute spot on the mesosternal tubercle, and 



WASP GENUS CERCERIS — SCULLEN 385 

the tegulae, which are light yellow; tegulae smooth and not elevated; 
enclosure minutely rugose transversely, with a prominent medial 
groove ; mesosternal tubercle present ; legs black or dark amber to the 
apical ends of the femora, beyond which they are yellow, again 
becoming darker on the tibiae ; hindfemora may be all black or show a 
lighter amber apical end; wings subhyaline except for the stigma, 
which is darker. 

Abdomen black except for a small patch on tergum 1, deeply 
emarginate band on terga 2 and 4, two lateral patches on tergum 3, 
which may be joined to each other by a narrow or broken line, solid 
narrow or broken band on tergum 5 ; pygidium with a broad rounded 
base narrowing to a much smaller rounded apical end, surface appear- 
ing velvet-like; venter dark amber, immaculate. 

Male. — Length 7 mm. Black and light yellow; deeply pitted; 
clothed with short silvery hairs. 

Head about one-fifth wider than the thorax, black except for large 
frontal eye patches, a patch on the medial lobe of the clypeus, base of 
mandibles, and scape, all of which are light yellow; clypeal border 
with three subequal denticles on the medial lobe ; clypeal surface con- 
vex; mandibles smooth; antennae normal in form, scape yellow, 
peduncle light amber, flagellum light amber below, darker above. 

Thorax black except for elongate patches on the pronotum, the 
scutellum, and tegulae, which are light yellow; tegulae smooth and 
not elevated; enclosure minutely rugose transversely, with a prominent 
medial groove; legs dark amber apically to the middle of the femora, 
light amber beyond, becoming darker on the tarsi; wings subhyaline 
except the stigma, which is amber. 

Abdomen black with small patch on tergum 1, broad band on 
tergum 2, two lateral patches connected by a broken line on tergum 3, 
narrow bands on terga 4 to 5, elongate patch on tergum 6, all of which 
are hght yellow; pygidium oval with both ends rounded and subequal, 
surface pitted; venter dark amber, immaculate. 

C. kennicottii Cresson is quite variable over its extensive range. 
It varies in size, shade of yellow, and extent of yellow markings. 
The yellow markings are more yellow in the Eastern States (C. kenni- 
cottii kennicottii Cresson) and more white in the southern and western 
specimens (0. kennicottii belai Scullen). In the Southeast, the female 
of C. kennicottii kennicottii Cresson is nearest to C blakei Cresson. 
They can be separated by the reddish amber of C. blakei, by the termi- 
nal border of the clypeal process, and by the difference in the tegulae. 
The male of C. kennicottii kennicottii Cresson is close to the male of 
C. finitima Jinitima Cresson. These usually can be separated by the 
elevated tegulae on C. jinitima finitima Cresson. In the West, the 
female of C. kenniccottii beali Scullen can be separated from C. con- 



386 



PROCEEDINGS OF THE NATIONAL MUSEUM 



vergens Viereck and Cockerell and other closely related species by the 
differences in the clypeal process. The relative size of the clypeal 
process on the female varies. The males are difficult or impossible 
to separate in some cases. 

Types. — The type male of C. kennicottii Cresson, from Louisiana 
(Robert Kennicott), is at the Philadelphia Academy of Natural 
Sciences, no. 1952. The type male of C. eriogoni Viereck and 
Cockerell, from Dripping Sprmgs, Organ Mts., N. Mex., at flowers 
of Eriogonum (Cockerell), is at the Philadelphia Academy of Natural 
Sciences, no. 10377. 

Distribution. — Although not taken in any large numbers, this 
species is recorded from most of the states east of the Rocky Moun- 
tains, from southern Canada and Massachusetts south to the Gulf of 
Mexico, and west to Arizona. It is much more common in the South 
Central States. 

Prey record. — None. 

Plant RECORD. — Acacias^). (Arizona), Cassiasp. (Nebraska, Texas), 
Ceanothus sp. (Virginia), Chaerophyllum tointurieri (Texas), Chamao- 
crista fasciculata (Nebraska), Cicuta sp. (Virginia), Cicuta maculata 
(Colorado, Virginia), cotton (Arizona, Texas), Daucus carota (North 
Carolina, Ohio, Virginia), Enpatorium sp. (Texas), Euphorbia bicolor 
(Texas), Euphorbia marginata (Missouri) , Helianthus tuberosa (Kansas), 
Melilotus alba (Ohio), Pastinaea saliva (Colorado), Solidago sp. (Colo- 
rado, Texas), Tamarix gallica (Texas), willow (Salix), (Texas). 




Figures 21, 22. — Localities of: 21, C. kennicottii kennicottii Cresson; 22, C. kennicottii 
beali Scullen. 

18b. Cerceris kennicottii beali, new subspecies 

Figure 22 

Female. — Length 8 mm. Black with creamy-yellow to white 
marldngs; otherwise, very close to C. kennicottii kennicottii Cresson. 



WASP GENUS CERCERIS — SCULLEN 387 

The female type shows two breaks in the band on tergum 2 similar to 
the female of C. convergens Viereck and Cockerell but the clypeal 
processes are very different. Other females taken at the same time 
and place show the above bands varying from a solid line to a series 
of evanescent spots. 

Male. — ^Length 6 to 7 mm. Black with creamy-yellow markings; 
otherwise, very close to C. kennicottii kennicottii Cresson. 

Types. — The holotype female and the allotype male, from Scotts- 
dale, Ariz., May 13, 1961 (R. S. Beal), taken in copula, are deposited 
at the U.S. National Museum, no. 66157. Paratypes are as follows: 

Arizona: 9, Amado, July 3, 1957, cotton (G. D. Butler); cf, Apache, Cochise 
Co., 5000 ft., Aug. 4, 1955 (R. R. Dreisbach) ; 9, d", Avra Valley, July 6, 1955, 
July 5, 1957, cotton (Butler-Werner) ; 9 , Camp Verde, Sept. 4, 1957 (T. R. Haig) ; 
9, Sd^cT, Canelo, July 30, 1956, May 18, 1957, June 21, 1958 (G. D. Butler); 
2 9 9, Casa Grande, May 26, 1955 (G. D. Butler) ; 9 , 2 cf cf, same locality, June 20, 
July 17, 18, 1956, alfalfa (C. WiUiams) ; 2 c^ cT, Continental, June 14, 1955, desert 
willow (G. D. Butler) ; 2 9 9 , Continental, July 27, 1956, alfalfa (C. Williams) ; 
4cf cf, Coolidge, May 29, 1955, Acacia sp. (G. D. Butler); d", Coohdge, July 24, 
1946 (IT. A. Scullen); 9, Eloy, Sept. 28, 1955, alfalfa (G. D. Butler); 9, Eloy, 
June 20, 1956, alfalfa (G. D. Butler); 9, Kansas Settlement, July 28, 1955 
(Butler- Werner) ; 9, Litchfield Park, June 11, 1954, alfalfa (G. D. Butler); 9, 
cT, Marana, July 6, 1956, July 5, 1957, mesquite, mustard (G. D. Butler); cf, 
Marana, July 27, 1956, cotton (C. Williams); 5cf cf, Marinette, July 6, 1950 
(R. H. Beamer); 2 9 9, Mesa, June 9, 1955, alfalfa (G. D. Butler); cT, Oak 
Creek Canyon, July 9, 1952 (R. H. Beamer and party); 9, Patagonia, May 21, 
1955, mustard (G. D. Butler); 9, Phoenix, Sept. 17, 1933 (R. H. Crandall); cf, 
Phoenix, Aug. 8, 1950 (R. S. Beal) ; cf , Phoenix, May 27, 1938 (Christenson) ; 
9, Phoenix, 1100 ft., Aug. 10, 1946 (H. A. Scullen); 9, Portal, June 10, 1961 
(H. A. Scullen) ; &, Portal, 5000 ft., Aug. 5, 1955 (R. R. Dreisbach) ; 9 , Queen 
Creek, July 13, 1956, cotton (C. Williams); 9, Safford, June 24, 1954 (G. D. 
Butler); 4 9 9, 4c?'cf, Scottsdale, June 12, 1961 (R. S. Beal); 9, 9 mi. E. Su- 
perior, July 23, 1956 (Butler-Gerhardt) ; 9 , Superior, Boyce Thompson Arbo- 
retum, July 23, 1955 (G. D. Butler); cf , Tempe, Aug. 20, 1956, alfalfa (C. Wil- 
Hams); 9, Vernon, July 25, 1956 (Butler-Gerhardt); 9, Yarnell to Prescott, 
July 23, 1942 (H. A. Scullen); 9, Yuma, July 31, 1956, vetch (C. Williams); 
2(^ <f, Yuma Valley, June 14, 1957, July 11, 1956 (G. D. Butler). California: 
cf, Blythe, Riverside Co., July 8, 1956 (L. A. Stange) ; 2d'<^, Calexico, Imperial 
Co., July 11, 1957, Sept. 23, 1957 (E. I. Schlinger); cf, Gordons Wells, Imperial 
Co., Aug. 29, 1957 (E. I. Schhnger) ; 9 , Ripley, Riverside Co., Aug. 16, 1946, Heli- 
anthus annuus (P. D. Hurd) ; 9 , 4cf cf , Seeley, July 17, 1940 (R. H. Beamer and 
party). New Mexico: cf , Rodeo, 4000 ft., Aug. 3, 1959 (II. E. Evans). Texas: 
2cf cf, Alpine, July 8, 1942 (H. A. Scullen); cf, Austin; cf, Bexar Co., 535 ft., 
April 1929, Gaillardia ambhjodom (H. B. Parks); cf, Brownsville, 1921 (J. C. 
Bridwell); cf, Brownsville, July 11, 1945, cotton; 2d'cf, Bro^vnsville, May 17, 
1952 (Michener and party) ; cf , Dallas, Dallas Co., Oct. 4, 1952, SoUdago altissitna 
(L. H. Shinner); cf, Davis Mts., July 9, 10, 1942 (E. C. Van Dyke); 2 9 9, Davis 
Mts., July 9, 1942 (H. A. Scullen) ; 9 , 2 cf cf , El Paso, July 11, 1917 (J. Bequaert) ; 
cf , Williams Co., Aug. 25, 1933 (J. E. Gillaspy). 



388 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Distribution. — Arizona, New Mexico, and adjoining areas. 

Prey record. — None. 

Plant record. — As indicated under paratypes. 

19. Cerceris krombeini, new species 

Figures 23, 122a,b,c, 

Female. — Length 10 mm. Black with yellow markings ; punctation 
coarse; pubescence very short. 

Head sUghtly wider than the thorax; black except large frontal 
eye patches, patch on the upper surface of the clypeal process, 
patch back of the eye, base of mandibles, and a patch on the scape, 
all of which are yellow; clypeal border with four denticles, the medial 
pair separated from each other by a deep depression and somewhat 
larger than the lateral pair; clypeal process with the free margins 
converging to a blunt point, below which is a pair of prominent 
denticles; mandibles with one distinct denticle flanked on each side 
by a low and indistinct denticle; antennae normal in form. 

Thorax black except for a divided band on the pronotum, two 
patches on the scutellum, the metanotum, a small patch on the 
pleuron, and the tegulae, all of which are yellow; tegulae slightly 
inflated and smooth; enclosure smooth except for a medial groove and 
deep pits in the lateral angles; mesosternal tubercles small but very 
acute; legs black on the basal segments to or just beyond the middle 
of the fore- and midfemora, beyond which they are yellow, the hind- 
femora entirely black or fuscous, the hindtibiae yellow basally, 
fuscous apically, and the hindtarsi fuscous; wings subhyaline, clouded 
apically, and the stigma bladk. 

Abdomen with a semidivided patch on tergum 1, a broad band 
with rectangular emargination on the basal end of tergum 2, broad 
but deeply emarginate bands on terga 3 and 4, a broad band on 
tergum 5; venter immaculate; pygidium with the sides convex and 
the ends subequal and rounded. 

Male. — Length 8 mm. Black with light yeUow markings; puncta- 
tion coarse ; pubescence very short. 

Head about one-sixth wider than the thorax; black except for 
the entire face, base of mandibles, and scape, all of which are hght 
yeUow; clypeal margin with three denticles on the medial lobe, the 
medial denticle slightly the largest; hair lobes subequal in width to 
the lateral clypeal lobes; surface of the medial clypeal lobe normally 
convex; mandibles with two very low denticles; antennae normal 
in form. 

Thorax black except for a divided band on the pronotum, two 
spots on the scutellum, the metanotum, and the tegulae, all of which 
are light yellow; tegulae distinctly elevated but smooth; enclosure 



WASP GENUS CERCERIS — SCULLEN 



389 



smooth except for a medial groove and a few deep lateral pits; meso- 
sternal tubercles absent; all legs black basally to near the apical ends 
of the femora, beyond which they are yellow except for a dark patch 
on the hind tibiae and the apical tarsal segments, which become 
darker; wings subhyahne but lightly clouded apically. 

Abdomen black except for a small semidivided patch on tergum 1, 
a broad band on tergum 2, broad but deeply emarginate bands on 
terga 3, 4, and 5, and a band on tergum 6, all of which are yellow; 
pygidium with sides convex and both ends rounded and subequal in 
width. 

The female of C. krombeini Scullen is very close in size, structure, 
and color pattern to the female of C. astarte Banks but it can be dis- 
tinguished by the differences in the clypeal denticles, the form of the 
pygidium, and the color of the stigma of the wing, which is very 
light in the latter and black in the former. 




Figures 23-25. — Localities of: 23, C. krombeini Scullen; 24, C. neahminax Scullen; 25, C. 
poculum Scullen. 



Types. — The type female and allotype male of C. krombeini Scullen, 
both from Continental, Ariz., Aug. 24, 1959 (K. V. Krombein), are 
in the U.S. National Museum, no. 66158. Paratypes are as follows: 

Arizona: 4??, Continental, Aug. 2-4, 1959 (K. V. Krombein) ; 2 99, Madera 
Canyon, Santa Rita Mts., July 31, 1958 (R. M. Bohart) ; 9, 10 mi. E. Nogales, 
Sept. 2, 1957 (T. R. Haig) ; 9, 13 mi. NW. Nogales, Sept. 3, 1957 (T. R. Haig). 

Distribution. — This species is known only from 10 specimens 
taken in a lunited area in southern Arizona. 
Prey record. — None. 
Plant record. — None. 



390 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

20. Cerceris neahminax, new species 

Figures 24, 123a,b,c 

Female. — Length 8 mm. Color black and creamy yellow; normally 
pitted ; clothed with short silvery hairs. 

Head slightly wider than the thorax, mandibles with two centrally 
located subequal denticles, black except for sides of face, spot on 
center of front, and the clypeus, which are creamy yellow; clypeal 
border slightly extended centrally without denticles; clypeal process 
in the form of a short, thin ridge with the border approximately 
straight, half the width of the border extension and with a dark amber 
border; antennae normal in form, scape creamy yellow, peduncle dark 
amber, flagellum light amber below and dark amber above. 

Thorax black except for two elongate patches on the pronotum, the 
metanotum, and the tegulae, which are creamy yellow; tegulae smooth 
and not elevated; enclosure smooth except for a central groove and 
pitted lateral borders; mesosternal tubercles absent; legs black except 
foretibiae, foretarsi, midtrochanter, midtibiae, midtarsi, hindtro- 
chanter, and basal ends of hindtibiae, which are creamy yellow; wings 
subhyaline except for a clouded area at the apex and the stigma, 
which is dark amber. 

Abdomen black except for a small elongate patch on tergum 1, 
narrow bands on terga 2 to 5, and lateral patches on sternites 3 and 
4, which are creamy yellow; pygidium suboval with the basal end 
narrowed to a point and the apical end rounded. 

Male. — Unknown. It is probably close to the male of C. 
acanthophila Cocker ell. 

C neahminax ScuUen is very close to C. acanthophila Cockerell, 
from which it may be separated by the following characters: 

acanthophila neahminax 

pygidium: pyriform oval 

mesosternal tubercle: small, black absent 

mid- and hindfemora: yellow apically black, immaculate 

mid- and hindtrochanter: black cream 

clypeal process: subequal in width to about half the width of 

clypeal border ex- the clypeal extension 

tension 

clypeal border extension: margin sinuate margin not sinuate 

Types. — The type female from Sante Fe, N. Mex., June 11, 1935 
(E. C. Van Dyke), is at the California Academy of Sciences. Paratypes 
are as follows: 

Arizona: 9, Ashfork, July 22, 1932 (R. H. Painter); 9, Chiricahua Mts., 
July 27, 1957 (D. J. and J. N. KnuU). New Mexico: 9, Mountain Park, 
June 27, 1940 (L. C. Kuitert). Utah: 9 , Fool Creek Pass, Aug. 17, 1938 (G. F. 
Knowlton, F. C. Ilermston). 



WASP GENUS CERCERIS — SCULLEN 391 

Distribution. — Very rare. Arizona, New Mexico, and Utah. 
Prey record. — None. 
Plant record. — None. 

21. Cerceris poculum, new species 

Figures 25, 124a,b,c 

Female. — Length 9 to 10 mm. Black and creamy yellow; normally 
pitted; clothed with silvery hairs. 

Head slightly wider than the thorax; black except for large frontal 
eye patches, the clypeus exclusive of the free border, patch on frons, 
all of which are creamy yellow; clypeal border with a broad extension 
at the base of the process; clypeal process broad and short with the 
sides strongly curved downward, black along the border; mandibles 
with two closely joined denticles subequal in size; antennae normal in 
form, scape and peduncle black, flageUum amber below, dark above. 

Thorax black except for two elongate patches on the pronotum, 
two spots on the scuteUum, the metanotum, and the tegulae, all of 
which are creamy yeUow; tegulae smooth and not elevated; enclosure 
faintly ridged transversely along the medial groove, sparsely pitted 
laterally; mesosternal tubercle absent; mid- and hindlegs black basally 
to near the apical ends of the femora, beyond which they are creamy 
yellow except for an elongate patch of dark amber on each tibia and 
the tarsi, which become light amber; hindlegs have coxae black, 
trochanters creamy yellow, femora black, tibiae yellow with a dark 
amber patch and tarsi amber; wings subhyaline except for the stigma, 
which is amber. 

Abdomen black except for creamy-yellow bands on terga 1 to 5, 
broadly emarginate on terga 2 to 5, two lateral patches on sternites 
3 and 4; pygidium long, broader, and abruptly pointed basally, 
gradually narrowing and rounded apically. 

AIale. — Unlaiown. 

Types. — The type female, taken 23 mi. NE. Douglas, Ariz., 4450 ft., 
Aug. 1, 1946 (H. A. Scullen), is in the U.S. National Museum, no. 
66159. Paratypes are as follows : 

Arizona: ?, Herford, Cochise Co. (W. M. Mann). Texas: ?, Alpine, 
July 8, 1942 (H. A. Scullen); ?, Davis Mts., Jefif Davis Co., June 27, 1942 
H. A. ScuUen). 

Distribution. — -Recorded only from southern Ai'izona and western 
Texas. 
Prey record. — None. 
Plant record. — None. 



392 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

22a. Cerceris rufinoda rufinoda Cresson 

Figures 27; 125a,b,c 

Cerceris rufinoda Cresson, 1865, p. 121. — Packard, 1866, p. 61. — Cresson, 1887, 
p. 286.— Schlotterer, 1887, p. 491.— Robertson, 1889b, p. 303.— Ashmead, 
1890, p. 32.— Bridwell, 1898, p. 209.— Ashmead, 1899, p. 295.— Viereck and 
Cockerell, 1904, p. 138.— Smith, H. S., 1908, p. 370.— Banks, 1912a, p. 26.— 
Stevens, 1917, p. 422.— Mickel, 1917b, p. 452.— Strandtmann, 1945, 
p. 311.— Scullen, 1951, p. 1010. 

Female. — Length 9 to 10 mm. Black and reddish amber with 
light yellow markings; deeply and closely pitted; clothed with very 
short silvery hairs. 

Head slightly wider than the thorax; black except for two large 
frontal eye patches, round spots back of the compound eyes, base of 
mandibles and small patch on scape, all of which are light yellow; 
medial lobe of the clypeal border divided into an upper and lower part, 
the lower part is sinuate with a distinct, medial, denticle-like process, 
laterad of which are two less distinct extensions ; the upper part of the 
clypeal border appears as two semidivided rounded processes (which 
appear to be on the lower part of the free surface, but closer examina- 
tion shows them to be embodied in the divided border as indicated 
above) ; mandibles with two subequal denticles medially located (not 
evident on worn mandibles); antennae normal in form, scape amber 
with a yellow patch, peduncle dark amber, flagellum light amber 
below, darker above. 

Thorax black with occasional infusions of red on the propodeum 
and marked with light yellow as follows: elongate patches on the 
pronotum, two spots on the scutellum, the metanotum, spot on the 
pleuron, and on the tegulae; tegulae somewhat elevated and pitted; 
enclosm'e deeply pitted and with a medial groove; mesosternal tubercles 
small; fore- and midlegs dark amber to near the apical ends of the 
femora, beyond which they are yellow, tm-ning to light amber apically ; 
posterior legs with coxae dark amber; trochanters, femora, and remain- 
ing parts varying shades of amber ; wings subhyaline except the stigma, 
which is darker. 

Abdomen black, except first tergum is reddish amber, tergum 2 with 
a broad yellow band emarginate (in some specimens the reddish amber 
coloration extends onto the anterior part of the second tergum), terga 
3 to 5 with narrow emarginate bands of yellow; venter dark amber, 
immaculate; pygidium narrowing basally to a small rounded point, 
apical end broad and subtruncate. 

Male. — Length 9 to 10 mm. Black and reddish amber with yellow 
markings; deeply and closely pitted; clothed with very short silvery 
hairs. 



WASP GENUS CERCERIS — SCULLEN 



393 



Head about one-fifth wider than the thorax, black except most of 
face below antennal scrobes, mmute spots back of compound eyes, 
mandibles except tips, and the scapes, all of which are yellow (in some 
specimens the lower clypeal border has considerable black) ; clypeal 
border with three black denticles on the medial lobe; hair lobes 
normal in width; clypeal surface convex; mandibles smooth except for 
a single slight elevation, variable in size; antennae normal in form, 
scape yellow; peduncle dark amber, flagellum light amber below, 
darker above. 




Figures 26, 27. — Localities of: 26, C. rufinoda rufinoda Cresson (females only); 27, C. 
rufinoda crucis Banks. 



Thorax black except for two elongate patches on the pronotum, 
two patches on the scutellum, the metanotum, patch on the pleuron, 
and the tegulae, all of which are yellow; tegulae elevated to a variable 
amount and very lightly pitted; enclosure transversely rugose and 
with a medial groove ; mesosternal tubercles absent ; legs dark amber 
to amber basally to near the apical end of femora, the remainder of 
leg parts yellow, becoming darker on the tarsi, amber patch on apical 
end of hindtibiae ; wings subhyaline, cloudy at apex. 

Abdomen black except first tergum, which is reddish amber with a 
semidivided yellow patch, tergum 2 with a broad yellow band (basal 
black may be replaced with amber) , terga 3 to 6 with narrower bands 
of yellow somewhat emarginate; venter reddish amber anteriorly, dark 
amber posteriorly; pygidium sides convex with both ends subtruncate. 

The extent of the reddish amber varies considerably in different 
specimens. Those with the red covering most of the propodeum were 
described as the variety C. rufinoda crucis by Viereck and Cockerell. 
The present writer has given this form the rating of a subspecies, but 
this may be open to question. 



394 PROCEEDINGS OF THE NATIONAL MUSEUM vol. no 

A shadow of uncertainty must remain in regard to the correct 
identification of the males of rufinoda Cresson and those of closely- 
related species (until some worker can find a positive method of 
separating these and other difficult males) for the following reasons: 
(1) After extended studies, the writer has so far found no satisfactory 
characters to separate the males of rufinoda Cresson from the males of 
echo Mickel or other unknown, closely related species. (2) The 
species rufinoda Cresson was described from a male taken in an area 
(Colorado) where the females of both echo Mickel and the female long 
accepted as, but not proven to be, the female of rufinoda Cresson have 
been recorded in numerous localities. (3) Finally, two other closely 
related species, irene Banks and argia Mickel, of which the males are 
inseparable without association with the females, also are recorded 
from Colorado although less commonly. All of the above females are 
readily separated. 

Type. — The type male of C. rufinoda Cresson, from the Rocky 
Mountains, Colorado Territory (Riding), is in the Philadelphia 
Academy of Natural Sciences, no. 1955. 

Distribution. — This widely distributed species is recorded mostly 
through the Central States of Kansas, Nebraska, South Dakota, and 
Colorado. The following records show the extremities and diversity 
of its range : 

Alberta: 29 9, Medicine Hat, Aug. 9, 1939 (E. H. Strickland). Arizona: 
cf, 11 mi. SW. Eloy, Pima Co., June 19, 1953 (T. R. Haig); 4d'd', 11 mi. NW. 
Ft. Huachuca, July 10, 1952 (Beamer, Liang, La Berge) ; cf, Santa Rita Mts., 
July 10, 1952 (Beamer, Liang, La Berge); cf, 8 mi. S. Wilcox, July 12, 1952 
(Beamer, Liang, La Berge). Arkansas: 9 , Ouachita Mts., 25 mi. N. Ft. Smith, 
Aug. 30, 1939 (E. C. Van Dyke). California: 9, Hallelujah Junction, Lassen 
Co., July 7, 1949 (P. D. Hurd). Idaho: 9 , Lewiston, July 18, 1925 (C. L. Fox). 
Illinois: 9, Carlinville, 1902 (Robertson); 9, White Heath, Aug. 7, 1915. 
Montana: 39 9. North Carolina: 49 9, June 14, 1927, July 16, 1926, 
Aug. 3, 1926, Sept. 8, 1921 (C. S. Brimley). North Dakota: 9 , Beach, Aug. 25, 
1923 (C. N. Ainslie). New Jersey: 9 , Melage, Sept. 15, 1907. New Mexico: 
9, 20 mi. E. Deming, 4000 ft., Aug. 2, 1946 (H. A. Scullen). Ohio: 29 9, 
Columbus, July 21, Aug. 2, 1941 (J. E. Gillaspy); 29 9, Franklin Co., July 17, 
1942 (R. W. Strandtmann) ; 9 , Madison Co., June 28, 1942 (R. W. Strandtmann). 
Oregon: 9, Ontario, 2150 ft., Aug. 2, 1929 (H. A. Scullen). Texas: 9, 
Belfrage; 9 , Donna, July 21, 1933 (J. W. Monk) ; 2 9 9 , Fedor; 9 , Ft. Hancock, 
July 9, 1917; 9 , Lee Co., June; 9 , Victoria, Victoria Co., July 25, 1917. Utah: 
9 , Lincoln, July 22, 1949 (G. F. Knowlton) ; cT, Tooele, July 22, 1952 (G. F. 
Knowlton). Virginia: 9, Clifton, Aug. 6, 1933 (J. C. Bridwell) ; 9, Falls 
Church, July 24, 1913 (Wm. Middleton). Wyoming: 9, Grey Bull, Aug. 16, 
1927 (H. H. Knight). 

Prey record. — Tychius picirostris (Fabricius), Madison Co., Ohio, 
June 29, 1942 (Strandtmann, 1945, p. 311); Smicronyx squalidus 
Casey, Columbus, Ohio. 

Plant record. — Aster sp. (Kansas), Ceanothus americanus (Illi- 



WASP GENUS CERCERIS — SCULLEN 395 

nois), Chamaecrista fasciculata (Nebraska), Daucus carota (Ohio), 
Euphorbia sp. (Nebraska), Euphorbia marginata (Kansas), Helianthus 
sp. (Nebraska), Sium circutaefolium (North Dakota), Solidago sp. 
(Kansas), Thelesperma gracile (Kansas). 

22b. Cerceris rufinoda crucis Viereck and Cockerell, new status 

Figure 27 

Cerceris rufinoda var. crucis Viereck and Cockerell, 1904, p. 139. — Smith, H. S., 
1908, 370.— Banks, 1912a, p. 26.— Stevens, 1917, p. 422.— Mickel, 1917b, 
p. 453.— ScuUen, 1951, p. 1010. 

Female. — Length 9 to 10 mm. Black and reddish amber with hght 
yellow markings; deeply and closely pitted; clothed with very short 
silvery hairs. 

Head as for C. rujinoda rufinoda Cresson. 

Thorax black except for the propodeum, which is reddish amber 
exclusive of the enclosure, which is black; yellow markings as follows: 
two elongate patches on the pronotum, broken band on the scutellum, 
the metanotum, patch on the pleuron, and the tegulae; tegulae eleva- 
ted and pitted; enclosure rugose; mesosternal tubercle small; legs 
amber; wings subhyaline, clouded at the apex. 

Abdomen black except tergum 1, which is reddish amber with a 
semidivided yellow patch; tergum 2 with a broad yellow band; terga 
3 to 5 with deeply emarginate yellow bands; venter amber anteriorly 
becoming darker posteriorly. 

M ALE . — Indistinguishable . 

This is being raised from a variety to a subspecies. 

Type. — The type female, from Las Cruces, N. Mex., Sept. 25, 
1895 (Cockerell), is at the Philadelphia Academy of Natural Sciences, 
no. 10393. 

Distribution. — Mostly in the Rocky Mountain area and occasion- 
ally west. Specimens are as follows : 

California: 9, Topaz Lake, Mono Co., July 17, 1951 (A. T. McClay). Col- 
orado: ?, Boulder Co., Sept. 10, 18, 1925 (C. P. Custer); ?, Crowley Co., 
Aug. 4, 1933; 9, Denver, Sept. 5, 1920; 9, White Rock, near Boulder, 5100 ft., 
Aug. 13, 1919; 9, Julesburg, Aug. 4, 1899; 9, Jumbo Rs. [sic], Crook [Logan Co.], 
Aug. 12, 1901 (Sandhouse); 9, LaJunta, 4100 ft., Aug. 12, 1920. Idaho: 9, 
Bruneau, Owyhee Co., July 22, 1952 (W. F. Barr) ; Hot Creek Falls, Owyhee Co., 
July 22, 1952 (W, F. Barr). Kansas: 9, McKinney Lake, Kearny Co., July 12, 
13, 1954 (H. E. and M. A. Evens); 9, Norton Co., 2270ft., Aug. 24, 1912 (F. X, 
Williams); 9, PhUlips Co., 1940 ft., Aug. 30, 1912 (F. X. Williams); 4 99, 
Riley Co., Aug. 30, Sept. 3, 4 (J. B. Norton); 9, Riley Co., Sept. (Marlatt). 
New Mexico: 9, Deming, 4000 ft., Aug. 2, 1946 (H. A. Scullen). Oregon: 
9, Ontario, 2150 ft., Aug. 2, 1929 (H. A. Scullen); 9, 14 mi. E. The Dalles, 
July 17, 1929 (H. A. Scullen). Texas: 9, Davis Mts., Jeff Davis Co., July 9, 
1942 (H. A. ScuUen); 9, Fabens, Oct. 17, 1943 (R. W. Strandtmann) ; 9, Fedor, 
Sept. 23, 1897 (Birkmann). Utah: 9, Goshen, Aug. 12, 1949 (George Bohart) ; 



396 PROCEEDINGS OF THE NATIONAL MUSEXJM vol. ue 

9, Myton, Duches Co., June 30, 1958 (J. W. MacSwain) ; 9, Provo (O. E. 
Johnson) ; 9, Tooele, Aug. 23, 1937 (L. L. Hanson). 

Prey record. — None. 

Plant record. — Croton neomexicanum (New Mexico), Euphorbia 
sp. (Kansas), Helianthus sp. (Idaho), Solidago sp. (Utah). 

23. Cerceris truncata Cameron 

Figures 28, 126a,b,c 
Cerceris truncata Cameron, 1890, p. 121. 

Female. — Length 10 mm. Black with creamy-yellow markings; 
closely and deeply pitted; clothed with short silvery hairs, longer 
on the propodeum. 

Head subequal in width to the thorax; black except for frontal 
eye patches, clypeal process, lower portion of the clypeus, narrow 
patch between the antennae, spots back of compound eyes, and base 
of mandibles, aU of which are very light yeUow; clypeal border with 
a broad emarginate extension mesad, laterad of which are two much 
smaller, irregular, denticle-like processes; clypeal process only slightly 
convex, about as broad as long, with the free border almost straight, 
smooth, rounded at the corners and amber; mandibles with two 
small subequal denticles; antennae dark amber except the flagellum, 
which is somewhat lighter below. 

Thorax black except for a broken band on the pronotum, two 
spots on the scutellum, the metanotum, two triangular patches on 
the enclosure, two triangular areas on the propodeum, patch on the 
tegulae, and small spot on the mesosternal tubercle, all of which are 
light yellow; tegulae minutely pitted and not elevated; enclosure 
minutely pitted and faintly rugose transversely; mesosternal tubercle 
prominent; legs dark amber except for elongate yellow areas on aU 
tibiae, the fore- and midtarsi; wings subhyaline except for a clouded 
area along the anterior half and the stigma, which is dark amber. 

Abdomen black except for a broad band on tergum 1, narrow band 
on terga 2, 4, and 5, lateral spots on tergum 3, sternite 1, and most 
of sternite 2, small lateral spots on sternites 3, 4, and 5, aU of which 
are light yellow; elevated area on the anterior part of sternite 3; 
pygidium somewhat pyi'iform with both ends rounded. 

Male. — Length 9 mm. Black with creamy-yellow marldngs; 
closely and deeply pitted; clothed with short silvery hairs; longer 
on the propodeum. 

Head about one-fifth wider than the thorax; black except for large 
frontal eye patches, a large central patch on the clypeus, and minute 
evanescent spots back of the compound eyes, all of which are creamy 
yellow; clypeal border with a broad extension on the median lobe, 



WASP GENUS CERCERIS — SCULLEN 



397 



which has a sinuate margin but no distinct denticles; mandibles 
smooth; antennae normal in form, dark amber except the lower 
surface of the flagellum, which is lighter. 

Thorax black except for a divided band on the pronotum, two spots 
on the scutellum, the metanotum, two spots on the enclosure, two 
triangular areas on the propodeum, small evanescent spot on the 
pleuron, and a patch on the tegula, all of which are light yellow; 
tegulae lightly pitted and not elevated; enclosure relatively smooth 
except for a central groove and a few deep pits along the lateral 
borders; mesosternal tubercle absent; legs black except for elongate 
areas on the sides of the tibiae and the basal parts of the tarsi; wings 
subhyaline except for a clouded area along the anterior half and the 
stigma, which is dark amber. 




Figures 28-30. — Localities of: 28, C. iruncata Cameron; 29, C. vanduzeei vanduzeet B&nks; 
30, C vanduzeei eburnea Scullen. 

Abdomen black except for a broad band on tergum 1, narrow bands 
on terga 2, 4, 5, and 6, small lateral spots on tergum 3, sternite 1, 
and small lateral spots on sternites 2 to 6, all of which are light 
yellow; pygidium oval with ends subequal and rounded; light yellow 
patches to the sides of the pygidium. 

The extent of the light markings varies in both sexes. Some 
specimens may show little or no yellow on the face, and the spots 
on the enclosure may disappear. 

C. truncata Cameron closely resembles C. convergens Viereck and 
Cockerell and C. kennicottii Cresson. In both cases the clypeal 
process is the best distinguishing character. 

Type. — The type female of C. truncata Cameron, from Temax, 
Yucatan, Mexico (Gaumer), is at the British Museum, no. 21.1,433. 

Distribution. — Southern Arizona, southern New Mexico, and 
southern Texas. 

r742-463— 65 5 



398 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

Prey record. — Algarohius prospis (LeConte), Mimosestes amicus 
(Horn), Mimosestes protractus (Horn), Neltumius arisonensis (Schaef- 
fer), all from Tucson, Ariz., in 1957-1958, by F. Werner (1960, pp. 
43-44), taken on Mexican palo verde (JParkinsonia) . 

Plant record. — Casually about oleander (Arizona). 

24a. Cerceris vanduzeei vanduzeei Banks 

Figures 29, 127a,b,c 

Cerceris vanduzeei Banks, 1917, p. 114; 1947, p. 32. — ScuUen, 1951, p. 1010; 

1960, p. 80. 
Cerceris cojnplanata Mickel, 1917a, p. 340. — Banks, 1947, p. 31. — Scullen, 1951, 

p. 1006. 

Female. — Length 8 to 9 mm. Black with yellow markings; 
closely and deeply pitted; clothed with very short silvery hairs. 

Head slightly wider than the thorax; black except for large frontal 
eye patches, most of the central lobe and side lobes of clypeus, spot 
back of compound eyes, basal half of mandibles, and patch on the 
scape, all of which are yellow; clypeal border showing five distinct 
denticle-like extensions; clypeal surface with a medially located low, 
blunt elevation, below which is a pair of erect black denticles; mandi- 
bles with two denticles, the more apical one somewhat the larger; 
antennae normal in form, scape with a yellow patch, peduncle amber, 
flagellum hght amber below, dark above. 

Thorax black except for two elongate patches on the pronotum, 
two spots on the scutellum, the metanotum, variable patch (which 
may be divided) on the propodeum, patches on the pleuron, and the 
tegulae, all of which are yellow; tegulae smooth and not elevated; 
enclosure heavily ridged at 45° angle to central groove; mesosternal 
tubercle small; legs black basally to near apical ends of femora, 
beyond which they are largely yellow except for patches of amber on 
the tibiae, tarsi becoming darker; forefemora with a pronounced 
depressed area on the posterior side; wings subhyahne except for the 
stigma, which is amber. 

Abdomen black with yellow emarginate bands on terga 1 to 5; 
bands on terga 2 to 5 broadly and deeply emarginate, band on tergum 
2 broader than others; venter dark amber, immaculate or showing 
very small yellow spots laterally on some sternites; pygidium suboval 
with both ends rounded. 

Male. — Indistinguishable. Probably it is being confused with 
closely related species. 

The entire face of the female may be black except for the frontal 
eye patches. Mandibular denticles may be worn off so that the 
mandibles appear smooth. The clypeal elevation and surface denticles 



WASP GENUS CERCERIS — SCULLEN 399 

are similar to those of C. conifrons Mickel, but the two species are 
very distinct in other respects. 

Types. — The type female, from San Diego, Calif., June (E. P. Van 
Duzee), is at the Museum of Comparative Zoology, Harvard, no. 
10030. The type female of C. complanata Mickel, taken at Auburn, 
Calif., Sept. 20, 1916 (L. Bruner), is at the University of Nebraska. 
The allotype male of C. complanata Mickel, taken at the same location 
and date, is also at the University of Nebraska. 

Distribution. — The subspecies C. vanduzeei vanduzeei Banks is 
recorded only from California. 

Prey record. — None. 

Plant record. — Chaparel (California), Croton californicus (Cali- 
fornia), Eriogonum fasciculatum (CaUfornia). 

24b. Cerceris vanduzeei eburnea, new subspecies 

Figure 30 

Female. — Length 9 to 10 mm. Black with white markings; 
closely and deeply pitted; clothed with very short silvery hairs. 

Head about one-fifth wider than the thorax; black except for large 
frontal eye patches, large spot on medial lobe of clypeus, small spot 
back of compound eyes, basal part of mandible, and patch on scape, 
all of which are white; clypeal border with five distinct denticle-like 
extensions on the medial section; clypeal surface with a centrally 
located low, blunt elevation, below which are two black denticles 
very close to the clypeal border; mandibles with one prominent 
denticle, basal of which is a very small evanescent denticle; antennae 
normal in form, scape with a creamy-white patch, peduncle amber, 
flagellum light amber below and darder above. 

Thorax black except for two elongate patches on the pronotum, 
two spots on the scutellum, the metanotum, patch on the pleuron, and 
patch on the tegulae, all of which are creamy white; tegulae smooth 
and not elevated; enclosure heavily and irregularly ridged at a 45° 
angle to the medial groove and deeply pitted laterally; mesosternal 
tubercle present; legs black to near the apical ends of the femora, 
beyond which they are creamy white except for patches of amber on 
the tibiae and the tarsi, which are darker; wings subhyaline, becoming 
cloudy at the apex, except the stigma, which is amber. 

Abdomen black with white bands on terga 1 to 5; bands on terga 
2 to 5 emarginate; band on tergum 2 broader than others; pygidium 
suboval with both ends rounded; venter dark amber, immaculate. 

Male. — Unknown. Probably it is being confused with closely 
related forms. 

The females show considerable variation in the amount of white on 
the face. It is common for all of the white on the face to disappear 



400 PROCEEDINGS OF THE NATIONAL MUSEUM tol. lie 

except the frontal eye patches, which may be greatly reduced. The 
smaller denticle on the mandibles often is not evident. 

The subspecies C. vanduzeei eburnea Scullen is similar in structure 
and color pattern to C. vanduzeei vanduzeei Banks except that the 
markings of the former are white while those of the latter are yellow. 

Types. — The type female, from North Powder, Union Co., Oreg., 
July 24, 1938, Solidago (H. A. Scullen), is deposited with the U.S. 
National Museum, no. 66160. Paratypes are as follows: 

British Columbia: 9, Vernon, July 18, 1947 (Hugh B. Leech). California: 
9 , Turlock, Stanislaus Co., June 3, 1955 (P. D. Hurd). Idaho: 2 9 9, Acequia, 
Minidoka Co., June 30, 1957 (W. F. Barr) ; 9 , Bliss, Gooding Co., July 23, 1955 
(J. E. Gillaspy) ; 9 , 2 mi. W. Gardens, Boise Co., July 3, 1956 (W. F. Barr) ; 
9 , Glenns Ferry, July 8, 1927; 9 , Hagerman, July 1, 1930; 4 9 9, Hot Springs, 
Owyhee Co., July 4, 1951, July 12, 1952, July 5, 1956 (W. F. Barr) ; 9 , Kiler, 
Sept. 7, 1923 (Carl D. Duncan); 29 9, Melba, Canyon Co., July 10, 18, 1957 
(H. W. Homan); 9, Mountain Home, 3138 ft., Sept. 1, 1952 (W. F. Barr); 
9 , 12 mi. NW. Regina, Ada Co., July 11, 1952 (W. F. Barr). Oregon: 29 9, 
5 mi. NW. Adrian, Malheur Co., June 25, 1958 (R. K. Eppley) ; 9 , Alvoid Desert, 
Andrews, Harney Co., 4200 ft., July 4, 1927 (H. A. Scullen) ; 3 9 9 , 36 mi. E. 
Baker, Baker Co., 2450 ft., July 31, 1929 (H, A. Scullen) ; 9 , Blitzen Valley, 
Harney Co., July 16, 1936 (S. Jewett, Jr.); 3 9 9 , 3 mi. S. Elgin, Union Co., 
Aug. 18, 1937 (Bolinger and Jewett); 69 9, Grants Pass, Josephine Co., Aug. 
7, 1950 (H. A. Scullen) ; 9 , Herford, Baker Co., 3650 ft., July 21, 1940 (H. A. 
Scullen) ,9,12 mi. W. Huntington, Baker Co., 2225 ft., July 22, 1936 (H. A. 
Scullen) ; 3 9 9 , 4 mi. E. Juntura, Malheur Co., 1440 ft., July 14, 1940 (H. A. 
Scullen); 29 9, Klamath Falls (Algoma), Klamath Co., Aug. 23, 1958 (Joe 
Schuh); 9, North Powder, Baker Co., July 24, 1938 (H. A. Scullen); 49 9, 
Ontario, Malheur Co., July 16, 1940, Solidago sp. (H. A. Scullen) ; 6 9 9 , Powder 
River, 26 mi. E. Baker, 3000 ft., Aug. 9, 1937 (Bolinger and Jewett) ; 9 , Prairie 
City, Grant Co., 3250 ft., Aug. 12, 1929 (H. A. Scullen) ; 9 , Rowena, Wasco Co., 
July 15, 1923 (Carl D. Duncan); 9, Umatilla, Umatilla Co., July 14, 1931 (J. 
Nottingham); 9, Unity, Baker Co., 5800 ft., July 21, 1940 (H. A. Scullen); 
9, Wilderville, Josephine Co., Aug. 4, 1949 (H. A. Scullen). Utah: 9, Forest 
Grove, Aug. 8, 1949 (G. F. Knowlton). Washington: 39 9, Stratford, Grant 
Co., July 4, 1920 (R. C. Shannon). 

Distribution. — Pacific Northwest. 

Prey record. — None. 

Plant record. — Achillea sp. (Idaho, Oregon), Chrysothamnus sp. 
(Idaho), Cleomella sp. (Idaho), Daucus carota (Idaho, Oregon), 
Eriogonum sp. (Idaho), Eriogonum elatum (Oregon), Helianthus sp. 
(Idaho), Melilotus sp. (Idaho), Solidago sp. (Oregon). 

Group II 

This group is distmguished by the following characters: (1) There 
is no prominent elevation on the surface of the medial lobe of the 
clypeus; however, two small papilliform structures are just above the 
clypeal border. (2) The second abdommal segment is much broader 
than the first, especially noticeable on the female. (3) The meso- 



WASP GENTJS CERCERIS — SCULLEN 401 

sternal tubercle is absent from both sexes. (4) The termmal segment 
of the male antenna is distorted slightly, (5) The males have 
relatively inconspicuous denticles on the clypeal border or none at all. 
(6) The male hair lobes are normal in width. (7) Buprestidae are 
used as prey." 

25a. Cerceris californica californica Cresson 

Figures 31, 128a,b,c 

Cerceris californica Cresson, 1865, p. 128. — Packard, 1866, pp. 60, 63. — Cresson, 
1887, p. 282.— Schletterer, 1887, p. 487.— Dalla Torre, 1897, p. 454.— Ashmead, 
1899, p. 295— Woodworth, 1913, p. 94.— Rohwer, 1917, p. 244.— Banks, 1947, 
p. 20.— ScuIIen, 1951, p. 1006.— Linsley and MacSwain, 1956, pp. 71-84.— 
Krombein, 1958a, p. 197.— Scullen, 1960, pp. 75-77. 

Cerceris ferruginior Viereck and Cockerell, 1904, p. 134. — Viereck, 1960b, p. 234. — 
Johnson and Ledig, 1918, p. 24.— Banks, 1947, p. 25.— Scullen, 1951, p. 1007. 

Cerceris garciana Viereck and Cockerell, 1904, p. 135. — Scullen, 1951, p. 1008. 

Cerceris populorum Viereck and Cockerell, 1904, p. 135. — Banks, 1947, p. 25. — 
Scullen, 1951, p. 1009. 

Cerceris cognata Mickel, 1916, p. 408. — Scullen, 1951, p. 1006. 

Cerceris denticularis Banks, 1917, p. 113; 1947, p. 20.— Scullen, 1951, p. 1007. 

Cerceris interjecta Banks, 1919, p. 84. — Scullen, 1951, p. 1009. 

Cerceris calodera Banks, 1947, pp. 22-23.— Scullen, 1951, p. 1006. 

Cerceris illota Banks, 1947, p. 23.— Scullen, 1951, p. 1008. 

Cerceris isolde Banks, 1947, p. 24. — Scullen, 1951, p. 1008. 

Cerceris californica californica Scullen, 1961, p. 46. 

Female. — Length 13 mm. Black with yellow markings, which 
may become amber in limited areas; closely and moderately pitted; 
clothed with short silvery hairs. 

Head subequal in width to the thorax; face, area back of eyes, two 
patches on vertex, base of mandibles, scape of antennae, all are yellow; 
clypeal margin slightly extended from the medial lobe, emarginate 
in the center; clypeal surface slightly convex; clypeal surface denticles 
barely evident; mandibles with two denticles, the more apical one 
much the smaller and undivided, the more basal one larger and with 
two distinct apices, and the more basad of these two apices much the 
smaller; antennae normal in form. 

Thorax black except the pronotum, tegulae, patch on pleuron, 
scutellum, metanotum, most of propodeum, and small spots on the 
enclosure, all of which are yellow with limited amber margins; tegulae 
not elevated, and relatively smooth except for many minute pits; 
enclosure finely rugose; mesosternal tubercle absent; legs yellow with 
some margins showing amber; wings subhyaline, cloudy in the 
anterior area. 



" Bridwell has reported an exception that occurs when buprestids are not 
available. See under C. fumipennis Say (p. 412). 



402 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Abdomen with broad bands more or less emarginate with black on 
anterior center, margins with some amber; venter largely yellow with 
limited amber; pygidium broad basally, tapering gradually to a 
narrow rounded end apically. 

Male. — Length 10 mm. Black to amber with yeUow markings; 
close and moderately deep pits; clothed with short silvery hairs. 

Head one-sixth wider than the thorax; face yellow below antennae 
except clypeal margin, which is dark amber; clypeal border slightly 
extended from the middle lobe with three very indistinct denticle-like 
extensions; clypeal surface slightly convex; mandibles without 
denticles; antennae normal in form except for a slight curvature of 
the apical segment. 

Thorax black to amber with yellow markings; divided yellow band 
on the pronotum; scutellum black or with light spots; metanotum 
yellow or with a broken yellow band; tegulae not elevated, relatively 
smooth, yellow; enclosure smooth except for minute pits and a few 
large pits in the lateral angles; mesosternal tubercles absent; pro- 
podeum black to reddish amber; legs black to light amber to the apical 
end of the femora, largely yellow beyond; wings subhyaline, clouded 
along anterior border and apex. 

Abdomen black with broad bands on terga 2 to 6, more or less 
emarginate on terga 3 to 6; tergvmi 1 black becoming reddish amber 
on some specimens; anterior half of tergum 2 becoming reddish on 
some specimens; venter black to more or less infused mth amber and 
yellow; pygidium with sides subparallel and the apical end rounded. 

Variation. — C. calvfornica Cresson varies in color from the black 
and yellow form of the Northwest and the Rocky Mountains, repre- 
sented by the type of the species, to the extremely light forms 
represented by C. arno Banks. In these lightest forms, the black has 
been replaced completely by very light amber to yellow. The latter 
are found mostly in the southwestern desert areas of California and 
Arizona. It is no doubt largely due to this extreme variation in 
color that so many species have been erected by former workers who 
have seen relatively few specimens representing limited areas. It is 
a matter of opinion which of them, if any, should be considered sub- 
species. The present author herein has designated the extreme 
lighter forms, which are more limited in thek distribution, as the sub- 
species C. calvfornica arno Banks. Cerceris calvfornica Cresson may 
be confused with related species found in the same area such as C. 
completa Banks and C. grandis Banks. These are best separated by 
the characters given in the key. 

Types. — The type male of C. calijornica Cresson, from California 
is at the Philadelphia Academy of Natural Sciences, no. 1953. The 
type male of C. jerruginior Viereck and CockereU, from Deming, 



WASP GENUS CERCERIS — SCXJLLEN 403 

N. Mex., is at the Philadelphia Academy of Natural Sciences, no. 
10378. The type male of C. garciana Viereck and Cockerell, from 
New Mexico, is at the Philadelphia Academy of Natm-al Sciences, 
no. 10380. The type male of C. populorum Viereck and Cockerell, 
from New Mexico, is at the Philadelphia Academy of Natural Sciences, 
no. 10385. The type female of C. cognata Mickel, from Worland, 
Wyo., July 10, 1911 (L. Bruner), is at the University of Nebraska. 
The type female and allotype male of C. denticularis Banks, from 
Umatilla, Oreg., June 1882 (Samuel Henshaw), are at the Museum 
of Comparative Zoology, Harvard, no. 10028. The type male of 
C. interjecta Banks, from Utah, is at the Museum of Comparative 
Zoology, Harvard, no. 13767. The type male of C. calodera Banks, 
from Jacumba, Calif., Aug. 12, 1917 (W. M. Wheeler), is at the 
Museum of Comparative Zoology, Harvard, no. 27622. The type 
male of C. illoia Banks, from Arizona, is at the Museum of Compara- 
tive Zoology, Harvard, no. 23541. The type male of C. isolde Banks, 
from Arizona, is at the Museum of Comparative Zoology, Harvard, 
no. 23540. 

Distribution. — C. californica californica Cresson, in its numerous 
color variations, ranges from Idaho and Utah on the east to the 
Pacific Ocean, and from British Columbia to southern and western 
Texas. Approximately 2000 specimens of this common species have 
been studied by the ^vriter. 

Prey record. — The only intensive studies of the nesting habits 
and prey of this species were published by Linsley and MacSwain 
(1956). As would be expected, the prey was species of the beetle 
family Buprestidae. 

The following species were found to be collected by C. californica 
californica Cresson at Tanbark Flat, San Dimas Experimental Forest, 
in the San Gabriel Mountains of Southern California: Acmaeodera 
acuta LeConte, A. adenostomae Cazier, A. angelica Fall, A. coquilletti 
Fall, A. dohrni Horn, A. dolorosa Fall, A.fenyesi Fall, A. gemina Horn, 
A. hepburni LeConte, A. holsteni White, A. jocosa Fall, A. nexa Fall, 
A. perlanosa Timberlake, A. plagiaticauda Horn, A. prorsa Fall, A. 
quadriseriata Fall, A. sinuata VanDyke, Agrilus angelicus Horn, A. 
hlandus Horn, A. politus (Say), Anthaxia aeneogaster Cast, Chsysoboth- 
ris deleta LeConte, C. liicana Horn, C. femorata (Oliver). 

Plant record. — Acacia sp. (Chihuahua, Mexico), A. augustissima 
(Arizona), A. greggii (Arizona, Texas), alfalfa (Arizona), Asclepias sp. 
(Arizona, New Mexico), A. erosa (California), A. mexicana (Cali- 
fornia), A. specinsa (Utah), Baccharis sp. (Texas), B. glutinosa 
(Arizona, California), Bebbia juncea (California), Cercidium texanus 
(Texas), Cleome lutea (Oregon), Chaenactis sp. (California), Chilopsis 
linearis, desert willow (Arizona), Chrysothamnus sp. (California), 



404 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Ooluhhrina texensis (Texas), Condalia sp. (Arizona), C. obtosifolia 
(Texas), Croton sp. (Arizona), C. californicus (California), Dalea 
emoryi (California), D. schottii (California), Daucus carota (Oregon), 
Dithyrea wislizeni (New Mexico), Eriogonum sp. (California), E. 
elatum (Oregon), E.fasciculatum (California), E. iwdatum (California), 
Grindelia sp. (California), Helianthus sp. (Oregon), hollyhock (Texas), 
Hyptis emoryi (California), Isocoma helerophylla (California), Lepidium 
medium (Arizona), Melilotus sp. (California), M. alba (Arizona, Idaho, 
Oregon), Monarda sp. (Texas), Mortonia sacbrella (Arizona), Pith- 
ecolobium sp. (Texas), Polygonum auberti (California), Prosopis sp. 
(California, Texas), Rhamnus crocea (California), Rhus sp. (Texas), 
Salsola pestijer (Utah), Sapidus saponaria (Arizona), Solanum 
elaeagnifolium (Arizona), Solidago sp. (Oregon), Tamarix sp. (Utah), 
Verbesina encelioides exauriculata (Arizona), water hemlock (Oregon), 
Wislizenia rejracta (Arizona), Yucca sp. (New Mexico). 




Figures 31-33. — Localities of: 31, C. californica californica Cresson; 32, C. calfornica arno 
Banks; 33, C. completa Banks. 



25b. Cerceris californica arno Banks, new status 

Figure 32 

Cerceris arno Banks, 1947, p. 19.— Scullen, 1951, p. 1005; 1960, pp. 75-76. 
Cerceris argyrotricha Rohwer, 1908, p. 324. — Scullen, 1951, p. 1005; 1960, pp. 
75-76. 

Female. — Structurally the subspecies C. c. arno Banks is like the 
typical form of californica Cresson. C c. arno Banks represents the 
extreme lighter form found in the southwestern desert area. In the 
lighter forms the black is more or less replaced by light amber or 
yellow. 

Head largely yellow bordered with amber ; vertex may become very 



WASP GENUS CERCERIS — SCULLEN 405 

dark amber in the ocellar area; the gena and occiput are mottled with 
yellow and amber. 

Thorax largely yellow and amber, which becomes mottled mth 
darker shades or becomes very dark amber on the dorsum of some 
specimens; scutellum, metanotum, and propodeum, including the 
enclosure, vary from all yellow to mottled with yellow and light amber; 
legs have variable amounts of yellow and light amber. 

Abdomen almost entirely yellow with some light amber margins. 

Male. — Males follow very much the color pattern and shades of 
the females. 

Head largely yellow with light amber areas about the ocelli and 
back of the eyes. 

Thorax mostly yellow with hght amber areas on the scutum, 
becoming very dark on some specimens; legs yellow and light amber. 

Abdomen largely yellow with light amber borders. 

Cerceris califomica Cresson is easily confused with related species 
found in the same areas, such as C. completa Banks and C. grandis 
Banks. These are best separated by the characters given in the key. 
The males of C. califomica arno Banks and C. grandis Banks especially 
are difficult to separate. The surface sculpturing of the enclosure 
has been found most useful by the author. The enclosure of the male 
grandis shows evidence of ridges which become prominent in the 
females of that species, while the enclosure of the male of califomica 
arno is relatively smooth except for a few pits in the lateral angles. 

Types. — The type female of C. arno Banks is at the Museum of 
Comparative Zoology, Harvard, no. 23541, and the type female of C. 
argyrotricha Rohwer is at the U.S. National Museum, no. 28485. 

Distribution. — Southwestern desert area of California and 
Arizona. 

Prey record. — None. 

Plant record. — Baccharis glutinosa (Arizona), cotton (Arizona), 
Croton (California), Eriogonum albertianum neomexicanum (Arizona), 
Prosopis juliflora (Arizona). 

26. Cerceris completa Banks 

Figures 33, 129a,b,c 
Cerceris completa Banks, 1919, p. 83; 1947, p. 17.— ScuUen, 1951, p. 1006. 

Female. — Length 15 mm. Black with yellow markings; normally 
pitted; clothed with short silvery hairs, which are somewhat longer 
in the region of the propodeum and the first abdominal segment. 

Head shghtly wider than the thorax; black except the entire face 
below the antennae other than the clypeal margin, base of the mandi- 
bles, the scape, two elongate patches on the vertex, and patch on the 



406 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iie 

genae bordering the compound eye, all of which are yellow; clypeal 
surface without elevations except two minute clypeal surface denticles 
just above the margin; clypeal margin with four denticle-like exten- 
sions, the lateral ones somewhat more acute than the two medial ones, 
which are definitely rounded; mandibles with two centrally located 
denticles, the basal one being the larger and more acute; antennae 
normal in form. 

Thorax black, except for two wide patches on the pronotum, small 
spots on the scutellum, the metanotum, patches on the propodeum, 
and patches on the pleuron, all of which are yellow; enclosure deeply 
ridged; mesosternal tubercle absent; legs yellow except for irregular 
patches on the dorsal sides of the femora and limited uregular areas 
on the coxae and trochanters; wings subhyaline. 

Abdomen: tergum 1 with lateral yellow patches; tergum 2 yellow, 
which all but surrounds a black area connected with the black an- 
terior border; terga 3, 4, and 5 deeply emarginate anteriorHy with a 
black area; tergum 6 yellow, exclusive of the pygidium, which is 
amber; venter largely yellow; pygidium pyriform with the apical end 
rounded. 

Male.— Length 11 to 15 mm. The type is smaller than most 
specimens examined by the author. Black with yellow markings; 
normaUy pitted; clothed with short silvery haks, which are somewhat 
longer m the region of the propodeum and the first abdominal segment. 

Head slightly wider than the thorax; black except the entire face 
below the antennae, large patch on the scape, and the base of the 
mandibles, all of which are yellow; clypeal surface slightly convex; 
clypeal border slightly extended from the medial portion and some- 
what sinuate; mandibles without denticles; antennae with apical 
segment sHghtly curved. 

Thorax black except widely separated patches on the pronotum, 
the tegulae, and a divided band on the scutellum, all of which are 
yellow; enclosure lightly ridged in the lateral angles; mesosternal 
tubercle absent; legs mostly black to near the middle of the femora, 
beyond which they are yellow; wings subhyaline but somewhat 
clouded along the anterior border; stigma light amber. 

Abdomen: tergum 1 black; tergum 2 yellow with a broad black 
patch in the center and a black anterior margm; terga 3 and 4 with 
yellow partly surrounding a broad black area in the center; terga 5 
and 6 largely yellow but deeply emarginate with black anteriorly; 
tergum 7 with yellow patches to the sides of the pygidium; venter 
largely yellow; pygidium with sides slightly convergmg to a truncate 
apical end. 

The female is here described for the first time. C. completa Banks 
closely resembles the darker specimens of C. californica Cresson, 



WASP GENUS CERCERIS — SCULLEN 407 

from which it can be separated by the form of the pygidimn, the 
sculpture of the enclosure, and the color pattern of the terga. 

Types. — -The type male of C. completa Banks, from Claremont, 
CaUf., is at the Museum of Comparative Zoology, Harvard, no. 13767- 

Distribution. — Scattered records througii California and one 
record from southern Oregon. Specimens are as follows: 

California: cf, Acton, Los Angeles Co., Aug. 25, 1958 (E. I. Schlinger) ; cf, 
Auburn, 1919 (L. Bruner) ; 2c?cr, Buttonwillow, Kern Co., June 17, 1957; cf, 
Cache Creek Canyon, Yolo Co., May 30, 1955 (E. I. Schlinger) ; cf, Davis, July 
11, 1948 (Brad Stevens) ; c?. Deep Spring, Inyo Co., July 16, 1953 (R. M. Bohart); 
cf. Deep Spring, July 16, 1953 (E. I. Schlinger); cf. Deep Spring, July 17, 1953 
(N. Malley) ; d". Deep Spring, July 17, 1953 (W. D. McLellan) ; 9 , Geyserville, 
July 31, 1927 (J. C. Bradley); cf, Gorman, Los Angeles Co., July 13, 1956, 
AscleTpias mexicana (E. G. Linsley) ; cf , Gorman, Los Angeles Co., July 13, 1956 
(P. D. Hurd); 2cfcf, Grass Lake, Siskiyou Co., 5000 ft., July 4, 1952 (M. 
Cazier and party); 2cf cf, Happy Camp, Siskiyou Co., July 8, 1958 (J. Powell); 
9 , Kern Co., northwest corner of county, on U.S. 41, June 20, 1952 (W. D. 
Murray); cf, Kernville, July 24, 1940 (L. S. Lipovsky); cf, 20 mi. S. Livermore, 
Alameda Co., June 6, 1959 (D. J. Burdick) ; cf , Lodoga, Colusa Co., July 12, 
1955 (E. A. Kurtz); 3cf cf, 3 mi. N. Lone Pine, Inyo Co., July 3, 1953 (J. W. 
MacSwain) ; cf , 2 mi. N. Lone Pine, Inyo Co., July 13, 1953 (W. D. McLellan) ; 
cf, Los Angeles Co.; cf, Los Gatos Creek, Mt. Diablo Range, Fresno Co., June 
6-8, 1907 (Bradley) ; cf , 15 mi. W. Mineral, June 16, 1941 (D. J. and J. N. KnuU) ; 
cf. Ridge Rute, Los Angeles Co., 3000-4000 ft., June 13, 1931 (Don Prentiss); 
cf. Ridge Rute, June 13, 1931 (H. A. Scullen); cf, 7 mi. S. San Antonio Ranger 
Station, Santa Clara Co., June 27, 1953 (R. O. Schuster); cf, Santa Monica Mts., 
July 9, 1912 (J. C. Bridwell); 2cf cf, Three Rivers, 600-800 ft., July 12-14, 1907. 
Oregon: 2cf cf. Gold HiU, July 12, 1930 (H. A. Scullen). 

Prey records. — None. 

Plant records. — Asclepias mexicana (California). 

27a. Cerceris dilatata dilatata Spinola 

Figures 34, 130a,b,c 

Cerceris dilatata Spinola, 1841, p. 118. — Dalla Torre, 1897, p. 457.— Scullen, 1961, 

p. 46; 1962, pp. 57-8. 
Cerceris maximiliani Saussure, 1867, pi. 1, p. 94. — Cameron, 1890, p. 111. — Dalla 

Torre, 1897, p. 467. 
Cerceris contracta Taschenberg, 1875, p. 396. — Schletterer, 1887, p. 489. — Dalla 

Torre, 1897, p. 456. 
Cerceris olymponis Strand, 1910, p. 140. 
Cerceris semiatra Banks, 1947, p. 25. — Scullen, 1951, p. 1005. 

Female. — Length 13 mm. Color black with brownish yellow 
markings; normal to closely pitted; clothed with short silvery hairs. 

Head slightly wider than the thorax, black except for narrow 
frontal eye patches and base of mandibles, all of which are brownish 
yellow, and the first three or four segments of the flagellum, which are 
light brown below; clypeal border is smooth, slightly extended, and 
emarginate in the center; clypeal surface is elevated slightly in the 



408 PROCEEDINGS OF THE NATIONAL MUSEUM vol. n« 

center but without a process; clypeal surface denticles almost indis- 
tinguishable; mandibles with two separated, centrally located denti- 
cles, the more apical one the smaller; antennae normal in form. 

Thorax black except for two small patches of brownish yellow on 
the pronotum; tegulae not elevated, with many minute pits and a few 
large pits ; enclosure surface with many minute pits and a few scattered 
large pits in the lateral angles; mesosternal tubercle absent; legs black 
except for small stripes on the tibiae and first tarsal segments of the 
first ajd second pairs of legs; wings subhyaline with a more clouded 
area along the anterior borders of the first pair. 

Abdomen black except for a band on the posterior border of the 
third tergum, wide emarginate bands on terga 4 and 5, patches laterad 
of the pygidium on tergum 6 and most of sternites 3, 4, and 5, all of 
which are brownish yellow; pygidium semioval with the apical end 
somewhat smaller. 

Male.^ — Length 9-10 mm. Black with brownish-yellow markings; 
deeply and closely pitted; clothed with short silvery hairs. 

Head slightly wider than the thorax, black except for elongate, 
narrow frontal eye patches; clypeal border slightly extended on the 
medial lobe, ending in three distinct, subequal denticles; hair lobes of 
the usual narrow type; mandibles without denticles; antennae largely 
black, terminal segments slightly curved. 

Thorax black except for two irregular small patches on the prono- 
tum; tegulae not elevated, with a few deep pits, between which are 
many minute pits; enclosure with an indistinct central groove, a few 
deep pits in the lateral angles and the general surface covered with 
scattered minute pits; mesosternal tubercles absent; legs black except 
for narrow stripes on the first two pair of tarsi; wings clouded along 
the anterior area. 

Abdomen black except for the apical half of tergum 3, most of terga 
4 to 7, and sternites 4 to 6, all of which are brownish yellow; pygidium 
with sides slightly convex and apical end truncate. 

Types. — A neotype male of C. dilatata Spinola, designated by the 
writer, is at the Instituto e Museo di Zoologia, University di Torino, 
Italy. The male and a female, both apparently determined by Spi- 
nola, are from Cayenne, French Guiana. A lectotype female of C. 
maximiliani Saussure, from Mexico, designated by the writer, is at 
the Museum Histoire Naturelle, Geneva, Switzerland. A lectotype 
female of C. covtracta Taschenberg, from Brazil, designated by the 
writer, is at the Zoologisches Institut, Martin-Luther-Universitat 
Halle (Saale), Germany. The holotype female of C. olymponis 
Strand, from Paraguay, is at the Zoologisches Museum, Humboldt 
Universitat, Berlin. The type male of C. semiatra Banks, from Pati- 



WASP GENUS CERCERIS — SCULLEN 



409 



gonia, Ariz., is at the Museum of Comparative Zoology, Harvard, 
no. 27620. 

Distribution. — This species has been taken from southern Arizona, 
through southern New Mexico, and southern Texas. It ranges south 
through Mexico and Central America and into South America. 

Prey record. — None. 

Plant record. — (AU from Arizona) Acacia sp., Baccharis glutinosa, 
Cephalanthus sp., Chilopsis linearis (desert willow), Cissus trifoliata, 
Condalia lycioides, Croton sp,, Erigeron canadensis, Eriogonum sp., 
Lepidium sp,, Melilotus alba, Prosopis sp. (mesquite), Sapindus 
saponaris, Wislizenia refracts (jackass clover). 




Figures 34, 35. — Localities of: 34, C. dilatata dilatata Spinola; 35, C. dilatata chisosensis 
ScuUen. 

27b. Cerceris dilatata chisosensis, new subspecies 

Figure 35 

Female. — Length 13 mm. Black with reddish-amber and brown- 
ish-yellow markings; structurally like C. dilatata dilatata Spinola but 
colors depart from the typical subspecies by having considerable 
reddish amber covering most of the propodeum exclusive of the 
enclosure, the first abdominal segment, and extending onto the second 
tergum. The legs are amber and not black. Color marking otherwise 
more extended as indicated below. 

Head black except for two large frontal eye patches and most of the 
medial clypeal lobe, base of mandibles, and the antennal scape, all of 
which are brownish yellow; clypeal margin and clypeal surface 
denticles are amber. 

Thorax black except for two elongate patches on the pronotum, 
the tegulae, and the metanotum, which are brownish yellow to amber, 
and the propodeum, which is largely reddish; enclosure black; legs 
amber; wings clouded with light amber. 



410 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Abdomen differs from that of dilatata dilatata Spinola by having the 
first abdominal segment reddish with the red extending irregularly 
onto the second tergum; the second tergum also has a broad irregular 
patch of yellow along the posterior border; venter largely yellow to 
amber; pygidium and remamder of tergum 6 is amber. 

Male. — Length 9-10 mm. Black with reddish-amber and brown- 
ish-yellow markings; structure like that of the subspecies dilatata 
dilatata Spinola. 

Head coloration typical of the subspecies dilatata dilatata Spinola 
except the antennae are somewhat lighter. 

Thorax coloration pattern follows very much that of the female 
except the red of the propodeum is somewhat less in extent; legs are 
light amber; wings are light amber. 

Abdominal coloration follows the pattern of the female of the 
subspecies. 

Types. — The type female and allotype male, from Chisos Mts., 
Big Bend National Park, July 6, 1942 (H. A. ScuUen), are in the U.S. 
National Museum, no. 66161. Paratypes are as follows: 

Arizona: ?, southern Arizona, August 1902 (F. H. Snow); 2cfcf, 5 mi. W. 
Southwestern Research Station, Portal, 5400 ft., July 18, 1956, Melilotus alba 
(M. Cazier) ; ? , same locahty, Aug. 24, 1956 (E. Ordway). Texas: 7 c? cf, Chisos 
Mts., Big Bend National Park (J. Bequaert); cf, same locality, July 6, 1942 
(E. C. Van Dyke); 7? 9, led^cT, same locality, July 3-6, 1942 (H. A. Scullen); 
cf, Davis Mts., Jeflf Davis Co., Aug. 22, 1936 (J. N. Knull); 9, cf, Davis Mts., 
July 10, 1942 (E. C. Van Dyke); lOd'd', Davis Mts., June 21, 29, July 9, 1942 
(H. A. Scullen); 2c?cf, Davis Mts., Madera Canyon Park, July 20, 1950 (Ray 
F. Smith); 9, El Paso, June 23, 1942 (E. C. Van Dyke). Mexico: 9, 
Chihuahua, Chih., Aug. 12, 1951, Baccharis glutinosa (P. D. Hurd) ; 2c?'c?, 18 
mi. W. Jimenez, Chih., Aug. 10, 1951, Baccharis sp. (H. E. Evans); cT, 33 mi. S. 
Hidalgo del Parral, 6400 f1., Oct. 24, 1957 (H. A. Scullen); 9, 10 mi. W. Nami- 
quipa, Chih., 6600 ft., July 3, 1947 (C. D. Michener) • 9 , San Juan Del Rio, 
Durango, 5200 ft., July 30, 1947 (C. D. Michener). 

Distribution. — This subspecies has been taken mostly in the 
Chisos Mountains of the Big Bend National Park, Texas, but limited 
numbers have been collected in nearby mountains and in northern 
Mexico. 

Prey record. — None. 

Plant record. — Baccharis glutinosa (Chihauhau, Mexico), Meli- 
lotus alba (Arizona). 

28. Cerceris fumipennis Say 

Figures 36, 131a,b,c 

Cerceris fumipennis Say, 1837, p. 381. — Dahlbom, 1845, p. 204. — Smith, F., 
1856, p. 465.— LeConte, 1883, p. 762.— Cresson, 1865, p. 113.— Packard, 
1866, p. 60.— Cresson, 1872, p. 227.— Tasehen berg, 1875, pp. 397-8.— 
Schletterer, 1887, p. 493.— Cresson, 1887, p. 282.— Marlatt, 1890, p. 147.— 



WASP GENUS CERCERIS — SCULLEN 411 

Robertson, 1892, p. 105.— Ashmead, 1894, p. 60.— Robertson, 1896, p. 
73.— Dalla Torre, 1897, p. 462.— Bridwell, 1898, p. 209.— Ashmead, 1899, 
p. 295.— Peckham, 1900, p. 90.— Smith, J. B., 1900, p. 519.— Viereck, 1903, 
p. 120.— Hartman, 1905, p. 66.— Smith, H. S., 1908, p. 366.— Smith, J. B., 
1910, p. 678.— Grossbeck, 1912, pp. 135, 299.— Banks, 1912a, p. 17.— 
Viereck, 1916, pp. 695, 696.— Mickel, 1917b, p. 447.— Britton, 1920, p. 
342.— Rau, 1922, pp. 30-31; 1928, p. 338.— Cartwright, 1931, pp. 269-270.— 
Scullen, 1951, p. 1008.— Krombein, 1952b, p. 95.— Linsley and MacSwain, 
1956, p. 74.— Evans, 1957, pp. 84-85, pi. 11.— Krombein, 1958b, pp. 101, 
110.— Evans, 1959, p. 156. 

Cerceris cincta Dahlbom, 1845, p. 204.— Smith, F., 1856, p. 438.— Cresson, 1887, 
p. 282. 

Cerceris unicincta Taschenberg, 1875, p. 397.— Patton, 1880, p. 403.— Cresson, 
1887, p. 282.— Schletterer, 1887, p. 505.— Dalla Torre, 1897, p. 462.— Ash- 
mead, 1899, p. 295. 

Female. — Length 15 mm. Black with very hmited creamy-yellow 
markings; deeply and closely pitted; clothed with short silvery hairs. 

Head subequal in width to thorax; black except for large patches 
mesad of the compound eyes, a patch on the medial lobe of the 
clypeus, and base of mandibles, all of which are light yellow; clypeal 
border with two widely separated denticles, between which is a 
broad extension emarginate in the center; clypeal surface denticles 
inconspicuous ; mandibles with two distinct, medially located denticles, 
the basal one much the larger; antennae normal in form, black. 

Thorax black except for two patches on the pronotum and the 
metanotum, which are creamy yellow; tegulae smooth except for a 
few scattered pits, not elevated; enclosure covered with minute pits, 
a few scattered lateral large pits, and a central groove ; mesosternal 
tubercles absent; legs black except for elongate blotchy patches on 
all tibiae; wings clouded. 

Abdomen black except for a broad band on tergite 2 and small 
lateral patches on tergite 3, which are creamy yellow, more or less 
blotched with black; venter black, immaculate; pygidium somewhat 
pyriform with the surface rugose. 

Male. — Length 10-12 mm. Black with creamy-yellow markings; 
normally pitted and clothed with short silvery hairs. 

Head shghtly wider than the thorax, black except for large eye 
patches on the face; clypeal border sMghtly extended, with a straight 
margin except for a slight medial denticle ; mandibles without distinct 
denticles; antennae black, normal in form. 

Thorax black except for a broken band on the prothorax and 
the metanotum, which are creamy yellow; tegulae lightly pitted and 
not elevated; enclosure with many minute pits and a few large pits; 
mesosternal tubercle absent; legs black except for creamy-yellow areas 
on all tibiae; wings clouded. 



412 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Abdomen black with a broad band on tergum 2 and lateral patches 
on terga 3, 4, and 5, which may be connected by a broken line along 
the posterior margins of the terga; venter black, pygidium slender 
and elongate with both ends blunt. 

Structurally, C. fumipennis Say is very close to C. califomica 
Cresson and C. dilatata Spinola, but the color patterns are quite 
different. 




f- IV 




1\; 



Figures 36-38. — Localities of: 36, C. fumipennis Say; 37, C. grandis grandis Banks; 38, 
C. grandis percna Scullen. 

Types. — Say's type male, from Indiana, apparently is lost. A 
neotype male, from 3 miles west of Kaskaskia River, Boulder, 111., 
June 21, 1938 (D. Bolinger), has been designated by the writer and 
is deposited at the U.S. National Museum. The type male of C. 
cincta Dahlbom, from "America," is at the Universitets Zoologiska 
Institution, Lund, Sweden. The type female of C. unicincta 
Taschenberg, from "Tenn.," should be at the Zoologisches Institut, 
Martin-Luther-Universitat, Halle (Saale), Germany. 

Distribution. — East of the Rocky Mountains to the Atlantic 
States and from southern Canada to the Gulf of Mexico. 

Prey record. — The prey-coUecting habits of C. fumipennis Say 
have been observed by more workers than any other species of Cer- 
ceris. With the exception reported below by Bridwell, the prey 
always has been some species of Buprestidae. The following quo- 
tation from unpublished notes by J. C. Bridwell, on file at the U.S. 
National Museum, is of special interest in this connection : 

This species belongs to a natural group in Cerceris, of which the species 
in many parts of the world all, so far as known, store their nests with adult 
Buprestidae. 

The season of flight for Buprestidae ends about August 1st but the wasps 
are then still active. The material below illustrates the substitute used 
when Buprestidae are no longer present. 



WASP GENUS CERCERIS — SCULLEN 413 

After the conclusion of the flight of the Buprestidae in 1933, the colony of 
Cerceris fumipennis under observation at Clifton, Virginia, stored their 
nests with adult beetles of the genus Chlamisus, Chrysomelid beetels resembUng 
Buprestidae only in their hard integuments and in their metalic luster. 

Other observers have found C. fumipennis Say collecting Bupresti- 
dae much later in the fall. No other observers have recorded the 
species collecting anything but Buprestidae. 

The following species of Buprestidae have been recorded or other- 
wise indicated as prey of C. fumipevnis Say: Acmeodera pulchella 
(Herbst) (Scullen, from Iowa), Actenodes acornis (Say) (Cartright, 
from S.C.), Agrilus arcuatus (Say) (Cartright, from S.C.), A. 
bilineatus (Weber) (Cartright, from S.C), A. abjectm Horn (Rau, 
from Kansas), A. ruficollis (Fabricius), Buprestis consularis Gory, 
B. fasciata Fabricius, (Cartright, from S.C), B. lineata Fabricius 
(Cartright, from S.C), B. macidipennis Gory (Cartright, from 
S.C), B. rufipes (OHver) (Cartright, from S.C), B. striata Fabricius 
(Cartright, from S.C), Chrysobothris azurea LeConte (Evans, from 
Ark.), C. blancliardi Horn (Cartright, from S.C), C. dentipes (Ger- 
mer) (Cartright, from S.C), C. femorata (Ohver) (Bridwell, from 
Kansas; Cartright, from S.C; Scullen, from Iowa), C. floricola 
Gory (Cartright, from S.C), C. lesueuri Castelnau (Cartright, 
from S.C), C. purpureovittata Horn (Evans, from Ark.), C. quadri- 
impressa Castelnau (Peckham, from Wis.), C. sexsignata (Say) (Cart- 
right, from S.C; Scullen, from Iowa), C. verdigripennis Frost (Cart- 
right, from S.C), Cinyra gracUipes (Melsheimer) (Evans, from Ark.), 
Conotrachelus sp., (Hartman, from Tex.), Dicerca caudata LeConte 
(Evans, from Ind.), D. americana (Herbst) (Cartright, from S.C), D. 
nuttalli var. consularis Gory (Cartright, from S.C), D. divaricata (Say) 
(ScuUen, from Iowa; Evans, from Ark), D. lurida Fabricius (Scullen, 
from Iowa; Krombein, from N.C; Cartright, from S.C), D. ob- 
scura (Fabricius) (Bridwell, from Kansas), D. punctulata (Schonherr) 
(Cartright, from S.C; Grossbeck, from Long Island, N.Y.; Robinson, 
from Va.), D. spreta (Gory), D. tuberculata (Castelnau) (Cartright, 
from S.C). 

Plant record. — Ammi majus (Texas), Ampelopsis arbor ea (Texas), 
Ceanothus sp. (North Carolina), C. americana (Massachusetts), C. 
canadensis (Massachusetts), Chinquapin sp. (Florida), Circuta 
maculata (Ohio), Cyrilla racemiflora (Florida), Daucus carota (Florida, 
Ohio), Erigeron quercijloris (Florida), hydrangia (North Carolina), 
Melilotus alba (Florida, Iowa, Kansas, Ohio, Texas), Monarda citrio- 
dera (Texas, New Jersey), tea (Florida), Polygonum hydropiperoides 
(Florida), Polytaenia nuttallii (Ohio), Ptillmaius capillaceum (Florida), 
Quercus virginiana (North Carolina), Rhus copallinum (North Caro- 
lina), Salix sp. (Texas), Solidago sp. (North Carolina), Spiraea telici- 

742-463—65 6 



414 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

folm (Massachusetts), Stillingia sylvalica (Texas), water parsnip 
(Texas), Vitex negundo (Texas). 

29a. Cerceris grandis grandis Banks 

Figures 37, 132a,b,c 

Cerceris grandis Banks, 1913b, p. 423; 1947, p. 10.— ScuUen, 1951, p. 1008. 

Female. — Length 19 mm. Light yellow amber with limited 
markings of medium amber; punctation sparse and shallow; 
pubescence very short except in the region of the propodeum and first 
abdominal segment. 

Head subequal in width to the thorax; Ught yellow amber except 
patch in ocellar area, small patch on occiput and clypeal margin, all 
of which are light amber, and the apical part of the mandibles, which 
are dark amber; clypeal margin with two acute denticles opposite the 
junction of the middle and lateral lobes of the clypeus, and two mesal 
denticles that are broad and flat; clypeal surface convex with two 
conspicuous small surface denticles just above the clypeal border; 
mandibles with one broad but pointed denticle, apicad of which is a 
depression; antennae normal in form with the scape mostly yellow 
and the remaining segments light amber. 

Thorax largely hght yellow amber with a broad mesal stripe and 
two narrow lateral stripes of amber on the mesotergum and more or 
less amber along all sutures; enclosure heavily ridged at 45° angle to 
the base; legs yellow; wings subhyaUne. 

Abdomen all yellow; pygidium with sides narrowing abruptly to a 
rounded apical point. 

Male. — Length 14 mm. Coloration close to that of the female 
but with somewhat more amber; punctation and pubescence as in the 
female. 

Head subequal in width to the thorax; largely yellow with dark 
amber stripes between the antennal scrobes and ocelli, and an ir- 
regular area of amber on the vertex; clypeal surface convex; clypeal 
margin extended on the medial lobe with the margin showing three 
inconspicuous denticle-like points; mandibles smooth; antennae 
normal except the apical segment, which is slightly curved. 

Thorax largely light yellow amber except the mesotergum, which 
is light amber; tegulae not elevated and relatively smooth; enclosure 
smooth except for inconspicuous ridges that appear in the lateral 
angles. 

Abdomen entirely yellow except for slight amber shading along 
some sutures; pygidium with sides converging slightly to a truncate 
apex. 

C. grandis grandis Banks is very close to C. calijornica arno Banks, 



WASP GENTJS CERCERIS — SCULLEN 415 

from which it may be distinguished by the characters given in the 
key. The males especially are difficult to separate, and the writer 
has not yet found characters that are dependable in separating them. 

Type. — The type female of C. grandis Banks, taken at Fort Yuma, 
Ariz., is in the American Museum of Natural History, New York, 
no. 21181. 

Distribution. — Southern California and adjoining areas. Speci- 
mens are as foUows: 

Arizona: d', Arlington, July 12, 1956, cotton (G. D. Butler); 9,cf, Roll, 
June 24, 1955 (G. D. Butler); cf, Wellton, July 12, 1956, cotton (G. D. Butler); 
cf, Yuma, July 1940 (K. D. Snyder); cf, Yuma Valley, June 14, 1957 (G. D. 
Butler). California: cf, Blythe, June 23, 1946 (W. F. Barr, R. F. Smith); 
cf, Coachella Valley, June 26, 1933; ? , Imperial Co., June 1911 (J. C. Bridwell); 
9, Indio, June 27, 1936 (E. S. Ross); cf, same locality, 1954 (G. E. Bohart); 
2(^0", 15 mi. S. Indio, June 27, 1936 (M. Cazier); ?, northern end of Salton 
Sea, May 25, 1940; cf, Thermal, Riverside Co., July 2, 1956, Melilotus alba 
(M. Wasbauer). 

Prey records. — None. 
Plant records. — None. 

29b. Cerceris grandis percna, new subspecies 

Figure 38 

Female. — Length 19 mm. Structurally like C. grandis grandis 
Banks in all respects except for color. The subspecies percna is 
black with yellow markings and a limited amount of amber. 

Head black or amber except face below antennal scrobies, base of 
mandibles, broken areas back of compound eyes, and two small 
patches on the vertex, all of which are yellow; en the genae outside of 
the above yellow areas back of the compound eyes are amber in the 
type but black on the paratypes; scape amber but remainder of 
antennae almost black. 

Thorax black except for a divided band on the pronotum, two 
small spots on the scutellum, the entire metanotum, two large areas 
on the propodeum, and a small patch on the mesopleuron, all of which 
are yellow bordered with amber; an amber area covers much of the 
mesosternum; legs largely amber; wings subhyaline. 

Abdomen largely yellow except for black patches in the center of 
all terga joining the basal border but becoming very small on the 6th 
tergum; venter largely yellow infused with amber. 

The type appears to be somewhat discolored with cyanide. The 
paratype from San Diego is almost entirely black and yellow with 
relatively little amber. 

Male. — Unknown . 



416 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Types. — The type female, taken at Mokelumne Hill, Calaveras Co., 
Calif., 1500 ft. elevation (F. E. Blaisdell), is at the U.S. National 
Museum, no. 66162. Para types are as follows: 

California: ?, Werner's, San Diego Co., Aug. 1, 1921; ?, Los Angeles Co. 

Distribution. — Southern California. 
Prey record. — None. 
Plant record. — None. 

Group III 

This group is distinguished by the following characters: (1) A 
distinct lamella (or lamellae) appears on the free margin of the medial 
clypeal process. (2) The extension of the medial clypeal lobe on the 
male is noticeably narrow, with three more or less distinct denticles 
on its margin. (3) Mesostemal tubercles are absent from both sexes 
or are very small. (4) The terminal segment of the male antenna is 
distorted slightly. (5) Chrysomelidae are used as prey to feed the 
young, as recorded for three species {compacta Cresson, flavofasciata 
H. S. Smith, and robertsonii Fox). 

30. Cerceris cochisi, new species 
Figure 39 

Female. — Length 14 mm. Amber and yellow with little or no 
black or dark amber, punctation and pubescence normal. Structurally 
near the female or O. compacta compacta Cresson except that specimens 
of the former are slightly larger. Enclosure somewhat smoother. 

Head yellow below the antennal scrobies, two yellow spots on the 
vertex, and yellow patches back of the compound eyes; small dark 
areas between the ocelli; basal two-thirds of mandibles yellow, apical 
third and denticles dark amber; scape yellow and amber, remaining 
segments of antennae amber becoming darker apically. 

Thorax amber except pronotum, scutellum, metanotum, and tegulae, 
which are yellow; legs light amber; wings shaded with amber. 

Abdomen yellow except for tergum 1, anterior half of tergum 2, 
most of first 2 stemites, and the sixth segment, all of which are light 
amber. 

Male. — Length 12 mm. Amber and yellow with little or no black 
or dark amber, punctation and pubescence normal. Structurally near 
the male of C. compacta Cresson, except that specimens average slightly 
larger. Enclosure somewhat smoother. 

Head yellow below the antennal scrobies and patches back of the 
compound eyes; small black ocellar area; basal half of mandibles 
yellow, apical half dark ; scape yellowish amber, basal half of antennae 
amber, apical half very dark. 



WASP GENUS CERCERIS — SCULLEN 



417 



Thorax dark amber, clouded with variable amounts of very dark 
amber; pronotum, scutellum, metauotum, and tegulae yellowish 
amber; legs light amber; wings lightly shaded with amber. 

Abdomen yellow except first and second segments, which are 
largely amber. 

Some males show considerably more dark amber on the thorax and 
anterioral abdomenal segments than the type. 




Figures 39, 40. — Localities of: 39, C. cochisi Scullen; 40, C. compacta compacta Cresson. 

Types. — The type female, Lordsburg to Silver City, N. Mex., over 
5000 ft., June 17, 1942 (H. A. Scullen), and the allotype male, 2 mi. 
NE, Portal, Ariz., June 5, 1961, Condalia lycioides (H. A. Scullen), 
both at the U.S. National Museum, no. 66163. Paratypes are as 
follows : 

Arizona: cf, 3 to 5 mi. SW. Apache, Aug. 8, 1959, Baccharis glutinosa (H. E. 
Evans); cf, Bill Williams Fork, August (F. H. Snow); 4 0^6^, Boyce Thompson 
Arboretum, Superior, July 23, 1955, Condalia, Acacia greggii (G. D. Butler); 
2cf'd', 3 mi. NW. Casabel, July 17, 1960, Prosopis juliflora (J. E. Gillaspy); 
<S*, Cedar Creek, 15 mi. W. Apache, June 21, 1957 ( Butler- Werner) ; c?, Congress 
Junction, July; &, Continental, Aug. 12, 1957 (G. D. Butler); 9 , 6 mi. E. 
Douglas, Cochise Co., Sept. 5, 1958 (M. A. Cazier); cT, 5 mi. W. Fort Apache, 
June 21, 1957 (Butler- Werner) ; cf, Fort Thomas, Graham Co., 2700 ft., Aug. 7, 
1946 (H. A. Scullen) ; 9 , Granite Dells, July 5, 1950, Nolina microcarpa (J. G, 
Rozen); cf, 15 mi. E. Pearce, July 10, 1955, Sapindus (Butler- Werner) ; c?, 
Phoenix, July 11, 1932; cf, Portal 4800 ft., Aug. 4, 1959 (H. E. Evans); 29 9, 
5 mi. E. Portal, Cochise Co., Oct. 14, 1955 (C. and M. Cazier); 3 9 9, 12cfd', 
2 mi. NE. Portal, Cochise Co., 4700 ft., July 2, 1961, Condalia lycioides (H. A. 
Scullen); cT, same locality, July 11, 1961, Acacia greggii (H. A. Scullen); 2 0^ cf, 
same locality, July 8, 11, 1961 (H. A. Scullen); 7cf cf, same locality, July 30, 
1959, Aug. 1, 3, 1959, Baccharis glutinosa (M. Statham); 9, Prescott, July 29 
(J. N. Knull); 2cf cf, Sabino Canyon, Santa Catalina Mts., Pima Co., June 29, 
1955, Condalia, Cephalanthus (Butler- Werner) ; cf , Sedonia, June 29, 1950 (J. G. 
Rozen); cf, Sedonia, June 29, 1950 (L. D. Beamer) ; 2cf cf, Seneca, Salt River 
Bridge, July 15, 1956, salt cedar (G. D. Butler); cf, southern Arizona (Bequaert); 



418 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

cf , Texas Pass, Dragoon Mts., July 19-21, 1917 (R. C. Shannon); cf", Tombstone, 
July 27, 1955 (Butler-Werner); ? , 5 mi. NW. Tombstone, Cochise Co., 4200 ft., 
July 25, 1937 (Rehn, Pate, Rehn) ; 5cf cf, Tucson (F, H. Snow); cf, Tucson, 
May 7, 1957 (G. D. Butler); d^, 9 mi. N. Tucson, July 25, 1946 (H. A. ScuUen); 
cf, 20 mi. NW. Tucson, July 6, 1955 (Butler- Werner) ; 9, 20 mi. E. Tucson, 
Oct. 26, 1958 (G. and A. Ferguson); cf , White Mts., June 19, 1950 (H. O. Wright). 
California: ?, Anza, Riverside Co., July 5, 1956 (R. M. Bohart); cf, Blythe, 
Riverside Co., July 30, 1947, Tamarix gallica (J. W. MacSwain) ; cf, Devils 
Canyon, Cochell Valley, May 7, 1932; cT, Experimental Farm, Imperial Co., 
June 1912 (J. C. Bridwell) ; &, Imperial Co., May 1911 (J. C. Bridwell); cf, 
Morango Valley, San Bernardino Co., Aug. 29, 1934 (C. D. Michener) ; cf, Winter- 
haven, Imperial Co., June 16, 1956 (G. R. Ferguson). New Mexico: 2cf cf, 
Alamogordo, July 7, 1902; cf, 10 mi. NE. Lordsburg, June 17, 1942 (E. C. Van 
Dyke); ?, Lordsburg to Silver City, 5000 ft., June 17, 1942 (H. A. Scullen); 
?, Rodeo, Hidalgo Co., July 29, 1959 (M. Statham). Texas: cf, Bexar Co., 
May 2, 1930 (H. B. Parks) ; ? , Davis Mts., Aug. 29, 1947 (D. J. and J. N. KnuU) ; 
cf , Davis Mts., July 6, 1936 (J. N. Knull) ; ? , Devils River, May 3, 1907 (F. C. 
Bishop); ? , El Paso, June 29, 1921 (Carl D. Duncan); cT, Galveston, June 1900; 
9, McNary, Hudspeth Co., June 24, 1942 (H. A, Scullen). Mexico: <^, Las 
Parras, Baja, October 1923 (W. M. Mann) ; c?, 20 mi. N. Mesquital, Baja, Sept. 27, 
1941 (Ross and Bohart); d", Mulege, Baja, May 14, 1921 (E. P. Van Duzee) ; 
cf, Nogales, Mar. 5, 1951; cf , 10 mi. E. San Ignacio, Baja, Sept. 30, 1941 (Ross 
and Bohart). 

Distribution. — Southern Arizona, southern New Mexico, south- 
western Texas and northern Mexico. 

Prey record. — None. 

Plant record. — Acacia greggii (Arizona), Baccharis glutinosa 
(Arizona), Cephalanthus sp. (Arizona), Condalia lycioides (Arizona), 
Nolina microcarpa (Arizona), Prosopis juliflora (Arizona), Sapindus 
sp. (Arizona). 

31. Cerceris compacta compacta Cresson 

Figures 40, 133a,b,c 

Cerceris compacta Cresson, 1865, p. 127. — Packard, 1866, p. 63. — Cresson, 1872, 
p. 230; 1887, p. 282.— Robertson, 1887, p. 201.— Schletter, 1887, p. 488.— 
Robertson, 1889, pp. 297-304; 1891, pp. 570-571; 1894a, pp. 439, 455, 464; 
1894b, p. 112.— Dalla Torre, 1897, p. 456.— Ashmead, 1899, p. 295.— Smith, 
J. B., 1900, p. 519; 1910, p. 519.— Banks, 1912a, p. 17; 1912b, p. 107.— 
Viereck, 1916, pp. 695, 696.— Mickel, 1917b, p. 448.— Scullen, 1951, p. 1006; 
1960, p. 77. 

Cerceris solidaginis Rohwer, 1908, p. 323. — Banks, 1947, p. 15. — Scullen, 1951, 
p. 1010. 

Cerceris belfragei Banks, 1917, p. 114. — Scullen, 1951, p. 1005. 

Female. — Length 10 mm. Black with yellow markings, normally 
pitted and clothed with short silvery hairs. 

Head subequal in width to thorax, black except center of face 
between antennal scrobes, clypeal process margin, small spots back 
of the eyes, and basal part of mandibles, all of which are yellow; 
clypeal margin extended on the medial portion, without denticles; 



WASP GENUS CERCERIS — SCULLEN 419 

clypeal process lunar shaped with two oval lamellae attached to the 
border; mandibles with two prominent denticles, between which is a 
much smaller but distinct denticle; antennae normal in form with the 
scape yellow and remaining segments amber. 

Thorax black except for a divided band on the pronotum, two 
large patches or a divided band on the scutellum, the metanotum, 
two patches on the propodeum, patch on the mesopleuron (on some), 
and the tegulae, all of which are yellow; enclosure ridged at a 45° 
angle to the base; tegulae not elevated and sparsely pitted; mesosternal 
tubercles absent; legs dark amber basally, becoming light amber on 
the femora and yellow to amber beyond; wings subhyaline, but 
lightly clouded along the anterior margins. 

Abdomen: tergum 1 with a small yellow medial patch, tergum 2 
with a broad band on the posterior half; terga 3, 4, and 5 with deeply 
emarginate bands quite variable in the amount of yellow; tergum 6 
with a trace of yellow to the sides of the pygidium; venter dark amber; 
pygidium broadly rounded basally with the sides narrowing to a 
smaller rounded apical end. 

Male. — Length 10 mm. Black with yellow markings; normally 
pitted; clothed with short silvery hairs. 

Head subequal to thorax in width, black except for entire face 
below antennal scrobes, minute spots back of eyes, basal part of 
mandibles, and scape, all of which are yellow; clypeal border dark 
amber with a very narrow extension slightly swollen in the center; 
hair lobes normal; clypeal surface slightly convex; mandibles with 
one large, acute denticle; antennae normal in form. 

Thorax black except a widely divided band on the pronotum, two 
small patches on the scutellum, band on the metanotum, occasional 
patches on the propodeum, occasional patches on the mesopleuron, and 
the tegulae, all of which are yellow; tegulae not elevated and nearly 
smooth; enclosure smooth except for a deep central groove with deep 
pits along the lateral borders; mesosternal tubercles absent; legs 
very dark on the coxae, becoming amber on the femora and light 
amber on the tarsi; wings subhyaline becoming lightly clouded along 
the anterior margins. 

Abdomen: tergum 1 with a small yellow patch; tergum 2 with a 
broad band along the posterior margin or largely covering the entire 
tergum; terga 3 to 6 with a more or less deeply emarginate band; 
venter dark amber; pygidium broadly rounded basally, tapering to a 
narrower rounded apical end. 

Specimens of both sexes show considerable variation in the extent 
of the yellow markings; those from the more northern and eastern 
areas are inclined to be darker. Those from the southwestern areas 
may have considerably more yellow. 



420 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

Types. — The type female of C. compacta Cresson, from Illinois, is 
at the Philadelphia Academy of Natm-al Sciences, no. 1940. The 
type male of C. solidaginis Rohwer, from Las Cruces, N. Mex., is at 
the U.S. National Museum, no. 28486. The type female of C. heljragi 
Banks, from Texas, is at the Museum of Comparative Zoology, 
Harvard, no. 10029. 

Distribution. — Throughout the eastern and central states and 
southwest to southern California. 

Prey record. — Colaspis brunnea (Fabricius), Chrysomelidae, Clif- 
ton, Va. (Bridwell). 

Plant record. — Alfalfa (California, Arizona), Ampelopsis arborea 
(Texas), Asclepias sp. (New Mexico, Texas), Asclepias verticillata 
(Iowa), Baccharis sp. (Arizona), Baccharis salicina (Texas), Bisora 
americana (Texas), Ceanothus sp. (Virginia, North Carolina), Chernoth 
americanus (Virginia, Illinois), Cicuta maculata (Ohio, Virginia), 
cotton (Texas, Louisiana, Arizona, New Mexico), Croton lindheimeri 
(Florida), Daucus carota (Ohio, Texas), Euphorbia marginata (Kansas), 
Helianthus annuus (California), Ligustrum sp. (Florida), Melilotus 
alba (Ohio, Illinois, Cahfornia, Utah, Arizona), parsnip (Texas), 
peach (Georgia), Polytaenia nuttallii (Texas), Pycnanthemum (North 
Carolina), Quercus virginiana (North Carolina), Rhus copallina 
(Florida), Solanum elaeagnifolium (Arizona), Solidago sp. (Texas, 
Kentucky), Tamarix gallica (Kansas, CaUfornia, Utah). 

32a. CercerisflavqfasciataflavofasciataH. S. Smith 
Figures 41, 134a,b,c 

Cerceris flavofasciata H. S. Smith, 1908, p. 364.— Smith, J. B., 1910, p. 678.— 
Mickel, 1916, p. 409; 1917b, p. 448.— Rau, 1922, p. 31.— Scullen, 1951, p. 
1007.— Krombein, 1953, pp. 114, 123, 124, 125, 134; 1959, p. 197. 

Cerceris natallenus Brimley, 1927, p. 238.— Scullen, 1951, p. 1007. 

Female. — Length 14 mm. Black with yellow markings, punctation 
normal, pubescence normal. 

Head slightly wider than the thorax; black except for large frontal 
eye patches, a small patch between the antennae, and the basal part 
of the mandibles, all of which are yellow; cljrpeal border with two 
widely separated denticles, between which the margin is depressed 
slightly but otherwise smooth; clypeal process is in the form of a very 
broad crescent-shaped elevation equal to about seven-eighths of the 
distance between the eyes; on the margin of the process there is 
a lamellae consisting of two distinct oval membranes; mandibles with 
three distinct denticles, the more apical one being very large, the 



WASP GENUS CERCERIS — SCULLEN 421 

basal one medium in size, and the middle one very small but very- 
acute; antennae normal in form. 

Thorax black except for a yellow band on the pronotum semi- 
divided at the center and two small patches on the scutellum, which 
are amber; metanotum black; tegulae smooth, amber, and not ele- 
vated; enclosure relatively smooth except for a slight medial groove; 
mesostemal tubercle present but small; legs black basaUy, dark 
amber on the femora, becoming lighter amber on the apical parts; 
wings somewhat more clouded than in most species. 

Abdomen entirely black in most specimens except for a broad 
yellow band on the posterior half of tergum 2; on some specimens 
small lateral yeUo w spots appear on terga 1 and 3 ; venter black ; 
pygidium oval with the apical end somewhat narrowed and both 
ends rounded. 

Male. — Length 10 mm. Black with yellow markings; punctation 
and pubescence normal. 

Head subequal in width to the thorax, black except large frontal 
eye patches, interantennal carina, variable amounts on the medial 
lobe of the clypeus, base of mandibles, and an evanescent spot on the 
scape, all of which are light yellow; clypeal extension on the medial 
lobe of the clypeus narrow, subequal in width to the length of the 
epistomal suture, and with three subequal small denticles; clypeal 
surface convex, mandibles with one denticle; antennae normal in form. 

Thorax black except for a broken band on the pronotum and divided 
band on the scutellum; tegulae amber, smooth, and not elevated; 
enclosure with a light medial groove, relatively smooth but with distinct 
pubescence laterally; legs amber, coxae and trochanters very dark; 
wings somewhat clouded. 

Abdomen with little or no yellow on tergum 1; a broad band on 
the posterior half of tergum 2; narrow yellow lines along posterior 
margins of terga 3 to 6; venter dark amber; pygidium oval but with 
the apical end truncate. 

Superficially, both the female and male of C. jlavojasciata Smith 
look much like C. fumipennis Say, but they are readily separated by 
the clypeal structure in each case. The female of C. fumipennis Say 
has no elevated process on the clypeus. The male of the latter has 
a very narrow extension on the medial lobe. 

Types. — The type female of C. jlavojasciata Smith is at the Uni- 
versity of Nebraska. The type female of C. natallenus Brimley is 
at the U.S. National Museum, no. 40792. 

Distribution. — Rare through the Eastern States from South 
Dakota, Nebraska, and Kansas to the Atlantic Coast States; from 
Minnesota south to North Carolina. Specimens are as follows: 



422 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 41-43. — Localities of: 41, C. flavofasciata flavofasciata H. S. Smith; 42, C. flavofas- 
ciata floridensij Banks; 43, C. mandibularis Patten. 

Arkansas: 9 > Hot Springs, Oct. 6 (Andreas Bolter). Illinois: 9 Hanna 
City, July 21, 1940. Iowa: 9, Sioux City, Aug. 10, 1917 (A. W. Lindsay). 
Kansas: 3 9 9, Baldwin Co., July 8, Aug. 1, 1906 (J. C. Bridwell) ; 9, Black 
Jack Creek, Pottawatomie Co., July 2, 1953 (Evans and party) ; 9 , Black Jack 
Creek, July 9, 1959 (H. E. and M. A. Evans); 9, Lawence, June 18, 1922 
(C. H. Curran) ; 9 , Marshall Co., June 30, 1950 (R. L. Fischer) ; 9 , Pottawato- 
mie Co., July 25, 1952 (H. E. Evans); 39 9, Riley Co., July 12, (J. B. Norton); 
29 9, Riley Co., July 23 (Popenoe); 9 , Saline Co., July 16, 1923, (L. C. Wood- 
ruflf). Minnesota: cf, Lake City, July 7, 1921 (A. A. Nichol). Nebraska: 9, 
Lincoln, July; cf. North Bend, June 30, 1915 (E. G. Anderson). Nebraska: 
Child's Point, Lincoln, Maskell, Monroe Canyon, Omaha, Rulo, and South Bend, 
June 30 to Aug. 20, 1917a, on flowers of Melilotus alba, Ceanothus americanus, 
and Amorpha canescens (all by Mickel). New Jersey: 9, Camden, July 12, 
1891 (Wm. J. Fox); 2 9 9, 5 mi. W. Lake Wood, Ocean Co., Sept. 6, 1954 (G. 
Ferguson); 9, Lebanon State Forest, Burlington Co., Aug. 19, 1958 (Evans, 
Beneway); 9, Lucaston, Sept. 9, 1906. North Carolina: 9, Bastic, July 7, 
1921 (T. B. Mitchell); 15,9 9 lOd'd', Kill Devil Hills, Dare Co., July 31, Aug. 
1, 3, 1952, July 26, 1955, Aug. 3, 4, 7, 8, 9, 1958 (K. V. Krombein) ; d", Kill Devil 
Hills, Aug. 7, 1958 (T. B. Mitchell). Oklahoma: 9 , Pawnee Co., July 12, 1932 
(Deonier, Pritchard). South Dakota: d^, Hot Springs, July 12, 1924 (R. C. 
Severin). Virginia: 9, Camp Peary, Sept. 7, 1943 (G. E. and R. M. Bohart). 
Maryland: 9, 30 mi. S. Washington, D.C., July 7, 1931 (J. C. Bridwell). 

Prey record. — Cryptocephalus guttulatus Oliver, C. notatus Fab- 
ricius, C. notatus fulvipennis Hald (all from Baldwin, Kansas). 
Krombein (1959, pp. 197-198), who studied the biology of this wasp 
at Kill Devil Hills, N.C., found Cerceris flavofasciata H. S. Smith 
collecting the following chrysomelids : Chlamisus sp. (probably plicata 
(Fabricius)), Cryptocephalus mutabilis Melsheimer, Cryptocephalus gut- 
tulatus Oliver, Cryptocephalus sp. (probably guadrimaculatus Say), 
Bassareus clathratus (Melsheimer), and Bassareus sp. (probably sellatus 
Suffrian). 

Plant record. — Amorpha canecens (Nebraska), Ceanothus ameri- 
canus (Nebraska), Melilotus alba (Nebraska). 



WASP GENUS CERCERIS — SCULLEN 423 

32b. Cerceris flavofasciata Jloridensis Banks (new status) 

Figure 42 

Cerceris austrina Fox (male only), 1893, p. 556.— Dalla Torre, 1897, p. 453.— 
Ashmead, 1899, p. 296.— ScuHen, 1951, p. 1005.— Krombein, 1954b, p. 
235; 1956, p. 234 (new synonymy). 

Cerceris Jloridensis Banks, 1915, p. 403. — Scullen, 1951, p. 1008. — Krombein, 
1954b, p. 235. 

Female. — Length 12 mm. Black with ferruginous markings; 
punctation and pubescence average for the genus. Structurally like 
C. flavofasciata flavofasciata H. S. Smith. 

Head reddish amber except for dark amber to black patches just 
above the antennal scrobes, the ocellar area, patches on the vertex 
bordering the eyes and the lower genal area, and the following parts, 
which are yellowish amber: large frontal eye patches, frons above the 
clypeal process, lateral lobes of the clypeus, and the basal part of the 
mandibles; apical part of mandibles and mandibular denticles dark 
amber; antennae ferruginous becoming dark apicaUy. 

Thorax black except the pronotum, scutellum, metanotum, most 
of the propodeum exclusive of the enclosure, patch on the pleuron, 
and the tegulae, all of which are ferruginous; legs reddish amber; 
wings clouded with amber. 

Abdomen: tergum 1 and 2 reddish amber with a narrow yellowish 
line on the posterior half of tergum 2 ; terga 3 to 6 black with a trace 
of dark amber on the anterior part of tergum 3; sternites 1, 2, and 3 
largely ferruginous. 

Male. — Length 10-12 mm. Black with dark ferruginous mark- 
ings showing some yellowing; punctation and pubescence average; 
structurally like the nominate subspecies male. 

Head black except for the face, which is light ferruginous medially, 
becoming darker on the clypeus ; genae and vertex largely ferruginous ; 
width of the clypeal extension on the medial lobe about one-half the 
length of the epistomal suture, with three subequal small denticles 
on the margin ; mandibles with one prominent acute denticle; antennae 
normal in form. 

Thorax black except for a broad band on the pronotum, scutellum, 
metanotum, and large patches on the propodeum, all of which are 
dark ferruginous; enclosure smooth except for a light medial groove; 
legs amber; wings clouded. 

Abdomen black except for the first two segments, which are dark 
ferruginous with a narrow yellowish band on the apical border of the 
second tergum. 

The males of several closely related species and subspecies in 
Florida and adjoining areas have been confused by earlier workers. 



424 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ue 

Only recently has the present writer been able to clear up some of 
this confusion. A study of type material in the British Museum, in 
this country, and field studies by Dr. Krombein have helped to as- 
sociate correctly some of the sexes concerned. In describing C. 
austrina, Fox evidently had the two sexes wrongly associated His 
type female of the latter species is accepted here as the type of 
austrina Fox. The present writer here considers C. austrina Fox a 
synonym of C. robertsonii robertsonii Fox. The form described by 
him as the male of C. austrina Fox has been found recently by Dr. 
Krombein to be the male of a ferruginous color form of C.flavofasciata 
H. S. Smith. As the name austrina is preoccupied, this color form is 
given the name C. flavofasciata floridensis Banks. Banks described 
a male of this form under the name oijloridensis. 

The male of C. rujopida F. Smith, which is very close to the male 
of C.flavofasciata floridensis Banks, frequently has been confused with 
it by the writer and other workers in earlier studies. 

Types. — The allotype male of C. austrina Fox from Florida is at 
the Philadelphia Academy of Natural Sciences, no. 4756. The type 
male of C. floridensis Banks, from Gulfport, Florida (Reynolds), is at 
the Museum of Comparative Zoology, no. 13765. 

Distribution. — Southeastern States, North Carolina to Florida 
and west to Alabama. Specimens are as follows: 

Alabama: 9 , Mobile, June 1949 (H. P. Loding). Florida: 2? 9,0?, Cocoa, 
July 1944 (G. E. Bohart) ; cf , Flagler Co., July 30, 1955 (H. V. Weems, Jr.) ; 9 , 
Gainesville, Alachua Co., April 16, 1952; 9 , 2cf d', Gainesville, April 29, May 3, 
5, 1919 (P. W. Fattig); 9, Gainesville, May 4, 1923 (Alexander, Walker); cf, 
Gainesville, May 11, 1928 (G. B. Merrill); &, Gainesville, May 13 (Debna, 
Weems) ; 2cf cf , Gainesville, Aug. 28, 1960 (P. M. Marsh) ; cf , Haines City, Sept. 
17, 1954 (H. V. Weems, Jr.) ; d", Hollywood, May 6, 1927 (S. Greenicher) ;8d'd', 
Inverness, 1892, 1918 (C. Robertson) ; 9 , Jacksonville, Oct. 6-9, 1941 (Howard 
E. Evans) ; 9 , cf , Lake Placid, Archbold Biological Station, March 28, 29, 1957 
(H. E. Evans); 4c? cf, Lake Placid, April 1, 1954, June 22, 25, 28, 1962 (K. V. 
Krombein); d', Levy Co., July 13, 1954 (H. V. Weems, Jr.); 9 , Newman's Lake, 
Alachua Co., Sept. 4-6, 1929 (T. H. Hubbell) ; 2 9 9 , Orlando, March (R. and 
G. Bohart); cf, Orlando, April 11, 1925 (O. C. McBride) ; 9, Ponce de Leon, 
July 13, 1934 (M. E. Griffith); d, Sanford, May 2, 1955 (H. E. and M. A. 
Evans); cf. Silver Springs, July 28, 1962 (F. Jacot-Guillarmod) ; 9, Tutusville, 
April 25, 1923; d, Wagner, April 10, 1941 (H. T. Fernald) ; d, Winter Park, 
May 27, 1942 (H. T. Fernald); 9, "Florida." Georgia: cf, Augusta, Rich- 
mond Co., May 30, 1959 (R. R. Snelling) ; 9 , Groveland, Cannoche River, July 
28, 1913 (J. C. Bradley); 39 9, d, "N. Georgia"; 29 9, Okefenokee Swamp, 
July 10, 1934, July 27, 1939 (R. H. Beamer) ; 9 , Spring Creek, Decatur Co., Aug. 
10, 1946 (P. W. Fattig); &, St. Simons Island, May 13, 1932 (P. W. Fattig); 
2cf cf, Tifton, May 30, 1946; 39 9, Videlia, Aug. 29, 31, 1937, Aug. 20, 1946 
(P. W. Fattig); 9, Waycross, July 16, 1916. North Carolina: 9, Burgaw, 
Aug. 19, 1948 (T. B. MitcheU) ; c?, Kill Devil Hills, June 30, 1950 (K. V. Krombein) 



WASP GENUS CERCERIS — SCULLEN 425 

Prey records. — None. 

Plant records. — Ceanothus amencanus (New Jersey Tea), Ilex 
opaca, mango trees, avocado (Florida). 

33. Cerceris mandibularis Patton 

Figures 43, 135a,b,c 

Cerceris mandibularis Patton, 1880, p. 403. — Cresson, 1887, p. 282. — Schletterer, 
1887, p. 496.— Dalla Torre, 1897, p. 467.— Ashmead, 1899, p. 295.— Banks, 
1912, p. 17.— Mickel, 1918, p. 447.— Cartwright, 1931, pp. 269-270.— 
ScuUen, 1951, p. 1009. 

Female. — Length 1 1 mm. Black with yellow markings, punctation 
somewhat widely separated and shallow, pubescence short and 
silvery. 

Head subequal in width to the thorax; black except for most of the 
face below the antennal scrobes, base of mandibles, and scape, all of 
which are yellow; lower clypeal area and margin dark amber; margin 
of medial lobe slightly extended with two lateral denticles, between 
which is a broad extension; clypeal process broad and lunar shaped 
with a short but broad uniform lamination attached to the margin; 
mandibles with a broad expansion on the inner edge, the margin of 
which shows three indistinct denticles; antennae normal in form. 

Thorax black except for two widely separated narrow patches on 
the prothorax, two small patches on the scutellum, and the tegulae, 
all of which are yellow; tegulae smooth and not elevated; enclosure 
with a central groove and ridged at a 45° angle; mesosternal tubercles 
absent; legs dark basally, gradually becoming light amber to yellow 
apically; wings subhyaline. 

Abdomen: tergum 1 with a small divided patch of yellow; tergum 2 
with a broad deeply emarginate band on the posterior half; terga 3, 4, 
and 5 with narrow emarginate bands on the posterior margin; tergum 6 
black; venter black; pygidium oval with the apical part somewhat 
narrowed and both ends rounded. 

Male. — Unknown. 

Type. — As the original type of Cerceris mandibularis Patton has 
apparently been lost, a neotype has been designated. Taken at 
Glenside, Pa., July 5, 1905 (G. M. Green), it is deposited at the U.S. 
National Museum. 

The neotype selected is 2 mm. longer than the indicated length in 
the original description of the species and has more yellow. Struc- 
turally it agrees with the original description and is representative of 
the species as recognized by later workers (Cresson and Banks). 
The male has never been identified. It is doubtless close to the males 
of related species, with which it probably has been confused. The 
female of mandibularis Patton closely resembles such related species as 



426 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii« 

rohertsonii Fox but the clypeal processes and their lamellae easily 
separate them. 

Distribution. — This species has been taken mostly in the Eastern 
States from New York south to Georgia and west to Iowa. Even 
though widely distributed in the East, collection records are relatively 
few. The following specimens have been recorded by the writer: 

Alabama: 9, Kushla. District of Columbia: ?, Washington, June 22, 
1951 (N. Banks). Georgia: ? , Tallulah Falls, June 19-25, 1909 (J. C. Bradley). 
Illinois: 9, Danville, August 31, 1924 (A.S.B.); 9, Prophetstown, July 7, 1925 
(T. H. Frison). Iowa: 9 , Ledges State Park, Boone Co., July 29, 1950 (Jean 
LaFoon); 9, Lyon Co., July 7, 1936 (H. E. Jaques) ; 2 99, Lyon Co., July 7, 

1936 (D. Milspaugh). Louisiana: 9 , Claiborne Co., May 22, 1931. Maryland: 
9, Baltimore, July 1914 (F. E. Blaisdell) ; 9, Baltimore, July 1909; 9, Thomas 
Road, near Cumberland, June 28, 1953 (L. M. Walker). New jersey: 9, 
Riverdale, July 24, 1909 (L. B. Woodruff). New york: 9, Bohemia, June 20, 

1937 (K. V. Krombein); 9, White Plains, June 29, 1918 (J. Bequaert). North 
Carolina: 9 , Mars Hill, July 6, 1928 (J. B. Mitchell). Ohio: 9 , Hocking Co., 
May 20 (D. J. and J. N. KnuU) ; 9 , S. Bloomingville, July 2, 1946 (U. N. Lanham). 
Pennsylvania: 9, Glenside, July 5, 1909 (G. M. Greene); 9, Leighgap, July 1, 
1897; 9, Pittsburg, June 15, 1911 (Hugo Kalil). Tennessee: 9, Tennessee 
City. Virginia: 9, Dunn Loring, July 28, 1951 (K. V. Krombein) ; 3 9 9, 
Falls Church, June 7, 27, July 6 (N. Banks). West Virginia: 9, Monongalia 
Co., June. 

Prey record. — The only information available relative to the 
prey collected by Cerceris mandibularis Patton is reported by Cart- 
wright (1931, p. 269), who reports the wasps collecting a grasshopper 
nymph. This recorded observation is open to question unless verified 
by later reports. 

Plant record. — None. 

34a. Cerceris rohertsonii rohertsonii Fox 

Figures 44, 136a,b,c 

Cerceris rohertsonii Fox, 1893 p. 55. — Robertson, 1894a, p. 464; 1894b, p. 112. — 
Ashmead, 1899, p. 296.— Smith, J. B., 1900, p. 519; 1910, p. 678.— Banks, 
1912a, p. 17; 1912b, p. 107.— Viereck, 1916, p. 695.— Britton, 1920, p. 342.— 
Krombein, 1949, p. 271; 1950, p. 148.— ScuUen, 1951, p. 1010.— Krombein, 
1952c, pp. 288-295; 1952b, p. 95; 1953, pp. 119-121, 122, 123; 1954b, p. 235; 

1955, p. 234.— Linsley and MacSwain, 1956, pp. 77, 79.— Evans, 1957, p. 
86, pi. 12.— Krombein, 1958a, p. 197.— Evans, 1959, pp. 156-7. 

Cerceris austrina Fox (female only), 12 1893, p. 556.— Dalla Torre, 1897, p. 453. — 
Ashmead, 1899, p. 296.— Scullen, 1951, p. 1005.— Krombein, 1954b, p. 235; 

1956, p. 234 (new synonymy). 

Cerceris pleuralis H. S. Smith, 1908, p. 366.— Mickel, 1918, p. 452. 

Female. — Length 12 to 13 mm. Black with yellow markings; 
punctation average; pubescence short. 



12 For a discussion of this species, see under C. f. floridensis Banks (p. 424). 



WASP GENUS CERCERIS — SCULLEN 427 

Head subequal in width to the thorax; black except most of the 
face, a small patch back of the eye, base of the mandible, and a patch 
on the scape, all of which are yellow; clypeal border with four exten- 
sions, the medial two in the form of roimded carina and the two 
lateral ones denticle-like; the clypeal surface elevation broad, uni- 
formally convex, and with a lamella on the free margin, the lamella 
more or less emarginate but not divided; mandibles with three denti- 
cles, the most apical one the largest and pointing apically; antennae 
normal in form. 

Thorax black except for a divided band on the pronotum, two oval 
patches on the scutellum, small spot on the pleuron, and spot on the 
tegulae, all of which are yellow; tegulae low and smooth; enclosure 
with a light medial groove and light ridges at about a 45° angle to the 
base; mesosternal tubercle small; legs yellow to fulvous with a darker 
area on the base of the femora and on the two basal segments. 

Abdomen with terga 1 and 6 immaculate, tergum 2 with a broad 
band on the posterior half, terga 3, 4, and 5 with narrow bands 
deeply emarginate; venter immaculate; pygidium oval with the apical 
end the narrower. 

Male. — Length 10 to 11 mm. Black with yellow markings; 
punctation average; pubescence short. 

Head subequal in width to the thorax; black except for the entire 
face, basal half of mandibles, and the scape, all of which are yellow; 
extension on the medial lobe of the clypeus narrow with three indis- 
tinct denticles; clypeal surface convex; hair lobes very narrow, less 
than the width of the lateral clypeal lobes; lateral clypeal lobes 
covered with prominent setae; mandibles with one small acute medial 
denticle; antennae with the apical segment slightly curved. 

Thorax black except for a divided band on the pronotum; a semi- 
divided band on the scutellum, spot on the pleuron, and a spot on the 
tegulae, all of which are yellow; tegulae low and smooth; enclosure 
without the usual medial groove, somewhat punctate laterally and 
lightly rugose along the anterior margin; mesosternal tubercle absent; 
fore- and midlegs with the first two segments black and yellow, the 
femora very dark basally, becoming light apically, remaining segments 
yellow; hindlegs with the coxae black and yellow, trochanters yellow, 
femora black basally, and the remaining segments largely amber with 
darker areas; wings subhyaline but clouded. 

Abdomen with terga 1 and 7 immaculate, tergum 2 with a broad 
band on the posterior half; terga 3 to 6 with narrow emarginate 
bands; venter with traces of yellow laterally on sternites 2, 3, and 4; 
pygidium with convex sides converging to a truncate apical end. 

Superficially, C. robertsonii robertsonii Fox closely resembles C. 
clypeata Dahlbom, C. atramontensis Banks, and other related forms in 



428 



PROCEEDINGS OF THE NATIONAL MUSEUM 



size and coloring, but the females can be separated easily by the form 
of the clypeal process, and the males, by the extension on the medial 
clypeal lobe. 

Types. — The type female of O. robertsonii Fox, from Smithville, 
S. Dak. (J. T. Aldrich), is at the Philadelphia Academy of Natural 
Sciences, no. 4755. The type female of C. austrina Fox, from southern 
Florida, is at the Philadelphia Academy of Natural Sciences, no. 4756. 
The type female, from Rock Co., Nebr., July 22, 1902, on Helianthus 
sp. (W. D. Pierce), and the allotype male, from Glen, Sioux Co., Nebr., 
4000 ft., Aug. 19, 1906 (P. R. Jones), of C. pleuralis H. S. Smith, are 
at the University of Nebraska. 

Distribution. — Throughout the Eastern States, from Nebraska 
and Kansas east to the Atlantic States and from southern Canada 
south to North Carolina, with one record from Texas. 

Prey record. — Rhabdopterus picipes (Ohver), Crypto cephalus 
notatus Fabricius, Pachybrachys dilatatus Suffrian. All are chrysom- 
elids, reported by Krombein (1953) from North Carolina. 

Plant record. — Melilotus alba (Kansas), parsnip (Minnesota), 
Quercus marilandica (North Carolina), Q. virginiana (North Carolina), 
Liguidambar styraciflua (North Carolina). 




Figures 44-46. — Localities of: 44, C robertsonii robertsonii Fox; 45, C. robertsonii bifidus 
ScuUen; 46, C. robertsonii emmiliosus ScuUen. 



34b. Cerceris robertsonii bifidus, new subspecies 

Figures 45, 137a,b,c 

Female. — Length 12 mm. Black with yellow and ferruginous 
markings; structurally like the nominate subspecies C. robertsonii 
robertsonii Fox except as indicated below. 

Head as in the nominate subspecies except that the clypeal lamella 
is divided completely to the base. 



WASP GENUS CERCERIS — SCULLEN 429 

Thorax like the nominate subspecies except that the yellow patches 
on the scutellum are fused lightly at the meson and the yellow marldngs 
on the legs replaced largely by ferruginous. 

Abdomen shows no important differences from the nominate 
subspecies. 

Male. — Unknown. 

Types.— The type female, from Kill Devil Hills, N. C, Aug. 4, 1952 
(K. V. Krombcin), is at the U.S. National Museum, no. 66165. 
Para types are as follows : 

Georgia: ?, Head River, July 24, 1936 (P. W. Fat tig). Indiana: ?, Mineral 
Springs, July 29, 1923 (Owen Bryant). North Carolina: ?, Aberdeen, July 
16, 1951 (Beamer, Weed, Price); 5 9 9, Kill Devil Hills, June 27, July 1, 11, 14, 
1950, Aug. 3, 1952 (K. V. Krombein) ; 9 , Raleigh, July 23, 1924 (C. S. Brimley). 
Virginia: 9, Princess Anne Co., Aug. 1, 1928. 

Prey record. — None. 
Plant record. — None. 

34c. Cerceris robertsonii emmiltosus Scullen 

Figure 46 
Cerceris robertsonii emmiltosus Scullen, 1964, p. 144. 

This subspecies is very similar to C. robertsonii miltosus except for 
the form of the lamella on the clypeal process. The lamella of 
emmiltosus has little or no emargination and is never completely 
divided at the meson as miltosus. Both subspecies are found in the 
Florida area. 

Type. — The type female of C. robertsonii emmiltosus Scullen, from 
Miami, Fla., March 29-30, 1953 (K. V. Krombein), is at the U.S. 
National Museum, no. 66166. 

DisTRiBUTiox. — Florida and Georgia. 

Prey record. — None. 

Plant record. — None. 

34d. Cerceris robertsonii miltosus, new subspecies 

Figure 47 

Female. — Length 11 mm. Black with j^ellow and ferruginous 
marldngs ; structurally^ like the nominate subspecies except as indicated 
below. 

Head black except the face, basal two-thirds of the mandibles, and 
the scape, all of which are yellow; clypeal lamella divided medially to 
the base. 

Thorax black except the entire pronotum, scutellum, metanotum, 
and the tegulae, all of which are ferruginous ; legs ferruginous, becom- 
ing more yelloAvish apically, wings clouded. 

742-463—65 7 



430 



PROCEEDINGS OF THE NATIONAL MUSEUM 



VOL. 118 



Abdomen with tergum 1 ferruginous, tergum 2 ferruginous except 
for a broad black area anteriorly, tergum 3 with a narrow band, which 
becomes broad lateraUy, tergum 4 with a broken Ime posteriorly, 
tergum 5 with lateral patches only; all abdominal markmgs become 
progressively more yellow posteriorly. 

Male.— Unknown (See note under C. f . jioridensi^ Banks, p. 424). 

This subspecies is very similar to C. robertsonii emmiltosus Scullen, 
from which it may be distinguished by the form of the clypeal lamella. 

Type— The type female, from Arcadia, Fla., April 23, 1953, on 
sand (K. V. Krombem), is at the U.S. National Museum, no. 66167. 
Paratypes are as follows: 

Florida: ? , Augustine; 9 , Cocoa, June 1944 (G. E. Bohart) ; ? , Jacksonville; 
9, St. Petersburg, July 1957 (G. Heinrich). 

Distribution. — Florida. 

Prey record. — None. 

Plant record. — None. 




Figures 47-50.— Localities of: 47, C. robertsonii miltosus Scullen; 48, C. rufopicta F. Smith; 
49, C. sandiegensis Scullen; 50, C. squamulijera Mickel. 



35. Cerceris rufopicta F. Smith 

Figures 48, 138 
Cerceris rufo-picta Smith, F., 1856, p. 467.-Cresson, 1865, p^ ISl^Packard, 
1866, p. 61.-Cresson, 1887, p. 282.-Scliletterer, 1887, p. 501.— Dalla Torre, 
1897^ p. 471.— Ashmead, 1899, p. 295.— Scullen, 1951, p. 1010. 

Female. — Unknown. 

Male.— Length 10 mm. Black with yellow and ferruginous 
markings; punctuation and pubescence average. 



WASP GENUS CERCERIS — SCULLEN 431 

Head subequal in width to the thorax; black except the entire face, 
basal two-thirds of mandibles, and a patch on the scape, all of which 
are yellow clouded with amber; extension of the medial clypeal lobe 
subequal in width to the length of the epistomal suture, with three low 
denticles; medial clypeal lobe convex; hair lobes narrow, covering 
slightly over half the lateral clypeal lobes; mandibles with one small 
denticle; antennae normal in form; flagellum ferruginous basally, 
becoming dark fuscous apically. 

Thorax black except for the entire pronotum, the scutellum, 
metanotum, tegulae, and patches on the propodeum, all of which are 
ferruginous; tegulae low and smooth; enclosure rugose with a deep 
medial groove and coarsely pitted laterally; mesosternal tubercle 
absent; legs ferrugiuous; wings subhyaline but clouded. 

Abdomen with the two anterior terga ferruginous with a yellowish 
line along the posterior border of tergum 2, terga 3 to 7 black with 
evanescent ferruginous lines along posterior margins of terga 3 and 4, 
becoming broader laterally, trace of ferruginous laterally on tergum 
5; venter black except the two anterior sternites, which are ferruginous, 
and a trace of ferruginous laterally on sternite 3 ; pygidium with sides 
convex and with ends subequal in width. 

For a discussion on the identification of the male of C. rufopicta, 
see note under C. flavofasciata floridensis Banks (p. 424). C. rufopicta 
F. Smith may prove to be the male of C. rohertsonii emmiltosus 
Scullen or C. rohertsonii miltosus Scullen. 

Distribution. — Florida and adjoining states north to North 
Carolina. Specimens are as follows: 

Florida: cf, Arcadia, May 11, 1961 (H. E. Evans); cT, Gainsville, May 24, 
1928 (G. B. Merrill) ; cf , Gainesville, May 17, 1959 (H. V. Weems, Jr.) ; &, 
Hillsboro Co., April 5, 1930 (C. L. Rabb) ; 40 d" cf , Inverness, 1892 (Robertson); 
2cf cf, Nov. 5 (Andreas Bolter); 2<^ d^, Juniper Springs, Marion Co., May 12-14, 
1954 (Karl V. Krombein) ; cf, Key Largo, April 27, 1946 (N. Mason); 26 c?" cf, 
April 1, 1954, June 22, 26, 1962 (Karl V. Krombein); 2cf cf, Lake Placid, 
Archbold Biological Station, April 15, 1947 (J. G. Needham) ; cf , same locality, 
April 6, 1956 (H. V. Weems, Jr.); 8cr cf, same locality, March 17, 25, April 1, 2, 
7, 1961 (S. W. Frost); cf, same locality, May 28, 1961 (H. E. and M. A. Evans); 
&, Miami, (P. W. Popenoe) ; c?, Miami, April 13, 1944 (D. C. Denning); cf, 10 
mi. SW. Ocala, Sept. 2, 1938 (Hubbell-Friauf) ; cf. Orange Co., April 12, 1930 
(N. W. Knowles); cf, Orlando, March 1944 (R. and G. Bohart) ; 4 d' cf , Sanford, 
May 2, 1955 (H. E. and M. A. Evans); cf, Tampa; cf, Wagner, April 10, 1941 
(H. T. Fernald); d', Welake, April 18-20, 1955 (H. E. and M. A. Evans); d'. 
Winter Park, April 29, 1936. Georgia: 2 rf" d", "Georgia" (Cresson) ; d', "Georgia" 
(Morrison); 4d^d^, "Georgia"; d^, "Georgia," February 1877. 

Prey records. — None. 

Plant records. — Ceanothus americanus (New Jeresy Tea), aovcado 
(Florida). 



432 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

36. Cerceris sandiegensis, new species 

FiGUEE 49 

Female. — Length 14 mm. Black with yellow markings; very 
little amber; punctation and pubescence normal; structurally near 
the female of C. compada compacta Cresson except it averages slightly 
longer and more slender. Enclosure somewhat smoother. 

Head black except face below antennal scrobies, two spots on the 
vertex, patch back of the compound eye, basal two-thirds of mandibles, 
and scape, all of which are yellow; apical third and denticles of 
mandibles very dark; peduncle and basal segments of flagellum 
amber, apical segments darker. 

Thorax black except band on pronotum, scutellum, narrow patch 
on metanotum, patch on the propodeum, tegulae, and spot on the 
pleuron, all of which are yellow; legs largely yellow with the femora 
becoming amber to dark amber dorsally; wings subhyaline with a 
more clouded area along the anterior part. 

Abdomen almost entirely yeUow except anterior part of terga 1 
and 2, sternite 1, and anterior part of sternite 2, all of which are very 
dark amber to black. 

Many females have the patch on the propodeum C-shaped and 
some are intermediate. 

Male. — Length 12 mm. Black with yellow markings; punctation 
and pubescence normal; structurally near the male of C. compacta 
compacta Cresson except it averages slightly larger; enclosure some- 
what smoother. 

Head black except for the face, two small spots back of the eyes, 
anterior two-thirds of mandibles, and scape, all of which are yellow; 
apical third of mandibles very dark; antennae beyond scape amber 
for the first 2 or 3 segments, gradually changing to a very dark amber. 

Thorax black except for a broken band on the pronotum, two 
small patches on the scutellum, patch on the propodeum, smaU patch 
on the pleuron, and the tegulae, all of which are yellow; enclosure 
relatively smooth; legs black and yellow; posterior trochanter largely 
yellow; all femora mostly dark amber to black; tibiae and tarsi 
mostly yellow, becoming amber on the apical tarsal segments; wings 
subhyaline with a more clouded area along the anterior part. 

Abdomen almost entirely yellow except the anterior part of terga 1 
and 2, sternite 1, and anterior part of sternite 2, all of which are dark 
amber to black. 

Types. — The type female, from two miles east of Anza, Riverside 
Co., Calif., July 14, 1956, on Oroton calif ornica (M. Wasbauer), and 
allotype male, IdyUwild, Riverside Co., Calif., June 27, 1956, on 



WASP GENTJS CERCERIS — SCULLEN 433 

Eriogonum fasciculatum var. polijolium (M. Wasbauer), both deposited 
at the California Academy of Sciences. Paratypes are as follows: 

Arizona: cf, Hereford, Cochise Co. (W. M. Mann); cT, Tempe, Aug. 4, 1917. 
California: 9, cT, Anza, Riverside Co., July 3, 1956 (L. A. Stange) ; ?, Anza, 
July 7, 1956, Eriogonum fasciculatum var. polifolium (E. G. Linsley) ; ?, Anza, 
July 14, 1956 (M. Wasbauer); 2 cf cT, 4 mi. N. Cajon, San Bernardino Co., June 
17, 1956 (G. R. Ferguson); 2 cfcf, 4 mi. N. Cajon Junction, San Bernardino 
Co., July 14, 1958 (E. I. Schlinger); d", same locality, July 1, 1958 (J. C. Hall); 
c?', Cajon Pass, San Bernardino Co., June 24, 1949 (H. E. Scott); cf, Cajon Pass, 
July 19, 1956 (H. R. Moffitt) ; d", Cuyamaca, July 17, 1949 (D. J. and J. N. 
Knull) ; 9 , Groveland, Aug. 20, 1953 (R. H. Goodwin) ; cf , Idylhvild, San Jacinto 
Mts., June 17, 1940 (E. C. Van Dyke); cf, Idyllwild, Aug. 3, 1935 (Jean Russell); 
9, cf, Idyllwild, June 19, 1951 (G. C. Bechtel) ; cf, Idyllwild, June 27, 1956, 
Eriogonum fasciculatum var. polifolium (M. Wasbauer); cf, Jacomba, June 19, 
1954 (D. J. and J. N. Knull); 3cf cf, Los Angeles (Coquillett) ; cf, Los Angeles, 
June 22, 1926 (Chas. H. Hicks); d', Newton, July 14, 1949 (D. J. and J. N. 
Knull); 9, cf, Oak Glen Lodge, San Bernardino Co., 5000 ft., July; 9, 4cf cf. 
Oak Grove, San Diego Co., June 6, 1940 (C. D. Michener) ; 9 9 9, 12cf cf, Pine 
VaUey, San Diego Co., Aug. 2, 1926, Aug. 1, June 27, 1927 (F. W. Kelsey) ; cf, 
Riverside, May 15, 1925 (Timberlake) ; 3 cf cf , Riverside, May 27 and June 1, 
1954 (J. C. Hall); 29 9, 7cf cf, San Jacinto Mts., July 1912, July 14, 1912 
(J. C. Bridwell) ; cf , San Jacinto IMts., Herkey Creek, June 20, 1940 (Fred H. 
Rindge) ; 9 , Tanbark Flat, Los Angeles Co., July 7, 1950 (H. F. Robinson) ; 9 , 
Temecula, Riverside Co., July 4, 1950 (J. W. MacSwain); cf, Warner Springs, 
San Diego Co., July 8, 1956, Eriogonum fasciculatum var. polifolium (P. D. Hurd); 
2cf cf, Warner Springs, Aug. 8, 1957, June 12, 1958 (H. R. Moffitt); cf, Warner 
Springs, July 4, 1956 (R. W. Bushing) ; cf , Warner Springs, June 12, 1958 (J. C. 
Hall); cf, Warner Springs, July 4, 1956 (R. M. Bohart) ; cf, 9 mi. S. Warner 
Springs, July 4, 1956 (R. M. Bohart); cf, Wrightwood, San Bernardino Co., 
5500 ft., July 1, 1956 (L. A. Stange); cf, Wrightwood, June 1, 1950 (H. R. 
Moffitt); 9, Yucaipa, San Bernardino Co., June 25, 1952 (L. D. Beamer and 
party). New mbxico: cf, Taos Co., June 14, 1956 (R. and K. Dreisbach). 

Distribution. — Southern CaUfornia, southern Arizona, and south- 
western New Mexico. 

Prey record. — None. 

Plant record. — Croton californica (CaUfornia), Eriogonum fas- 
ciculatum var. polifolium (CaUfornia). 

37. Cerceris squamulifera Mickel 

Figures 50, 139a, b,c 
Cerceris squamulifera Mickel, 1916, p. 411; 1917b, p. 451. — ScuUen, 1951, p. 1010. 

Female. — Length 10 mm. Black with yellow and amber markings; 
normal punctation, normal pubescence. 

Head subequal in width to the thorax; black except for the face 
below the antennal scrobes, large patches back of the compound 
eyes, the basal two-thirds of the mandibles, and the scape, all of 
which are yeUow; clypeal border slightly extended on the medial 



434 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lie 

lobe sinuate but without distinct denticles; clypeal process low and 
with the free margin equal to about one-third of the distance between 
the eyes; the clypeal lamellae not divided but slightly emarginate on 
the free margin and bent to about a 45° angle from the plane of the 
process surface; mandibles with three denticles, the more apical one 
very large, the medial one very small, and basal ones medium in size ; 
antennae normal in form. 

Thorax black except for a broad band on the pronotum, the scutel- 
lum, a narrow patch on the metanotum, a large comma-shaped 
area on the propodeum, and the tegulae, all of which are yellow; 
tegulae smooth and not elevated; enclosure smooth except for minute 
punctation and marginal deep pits; legs largely Ught amber, be- 
coming yellow on the more apical parts of the first and second pair; 
wings subhyaline. 

Abdomen largely yellow; first and second terga with the anterior 
one-third amber and posterior two-thirds yellow; terga 3, 4, and 5 
with broad yellow bands sUghtly emarginate, with black anteriorly; 
tergum 6 very dark amber; venter amber except for lateral yellow 
patches on sternites 3 and 4; pygidium oval, narrowing apically. 

Male. — Unknown. 

This species is very close to C. robertsonii Fox, but they can be 
separated easily by the form of the clypeal process. 

Type. — The type female of C. squamulijera Mickel from Imperial, 
Nebr., is at the University of Nebraska. 

Distribution. — Through the Western-Central States, from Ne- 
braska south through Kansas, Arkansas, Oklahoma, and into Texas. 
Specimens are as follows: 

Arkansas: 9, Arkansas River, June 8, 1956 (H. E. Evans, E. G. Matthews)- 
Kansas: 3 9 9, Hamilton Co. (F. H. Snow); 9, Meade Co., August 14, 1945 
(R. H. Beamer); 29 9, Medora, July 7, 1953 (Evans, Lin and Vashimoto); 9, 
Morton Co., (F. H. Snow); 9, Seward Co., August 16, 1911 (F. X. Williams); 
9, Stafford Co., June 30, 1934 (R. H. Painter). Nebraska: 39 9, Wallace 
Co., July 7, 1931, July 2, 3, 1933 (R. Roberts). North Carolina: 9, Southern 
Pines, June 13, 1918. Oklahoma: 9 , Alfalfa Co., August 8, 1932 (C. C. Dennier). 
Texas: 9, Clarendon Co., August 11, 1905 (Hunter); 9, Cypress Mills; 9, 
Galveston, Galveston Co.; 9, Galveston, May (F. H. Snow); 9, Galveston, 
June 24, 1924 (Trotter); 3 9 9, Galveston, June 1900. 

Prey record. — None. 
Plant record. — None. 

Group IV 

This group is distinguished by the following characters: (1) The 
females have a prominent cone-shaped process on the medial clypeal 
lobe, which is flanked on each side by a much smaller process attached 
to the lateral clypeal lobe. (2) The males have the medial denticle 



WASP GENUS CERCERIS — SCULLEN 435 

on the clypeal margin bicuspidate. (3) The pygidium of the male 
is very broad, ahnost oval. (4) The hair lobes of the male are very 
broad, almost meeting medially. (5) Tenebrionidae are used as 
prey by both species. 

38a. Cerceris femurrubrum femurrubrum Viereck and Cockerell 

Figures 51, 53, 140a,b,c,d,e,f 

Cerceris femurrubrum Viereck and Cockerell, 1904, p. 135. — Viereck, 1906b, 
p. 234.— Mickel, 1917b, p. 451.— Banks, 1947, p. 15.— ScuUen, 1951, p. 
1007; 1960, p. 78. 

Cerceris thione Banks, 1947, pp. 18-19. — ScuUen, 1951, p. 1007. 

Female. — Length 12 mm. Amber with light yellow markings, 
very Uttle black; structurally Uke C. Jemurrubrum athene Banks in 
all respects, except for color. 

Head all but totally amber except for a variable amount of yellow 
on the face mesad of the eyes. 

Thorax all amber to dark amber except for variable amounts of 
yellow on the prothorax, the scutellum, the metanotum, the propo- 
deum, and the pleuron. 

Abdomen amber with subequal emargmate bands on the terga 
along the posterior margins, aU of which are yellow; venter amber. 

Male. — Length 11 mm. Black with creamy-white markings and 
some amber parts; punctation and pubescence average. 

Head sUghtly \vider than the thorax; black except for the face, 
basal half of mandibles, very small spot back of the eye, and the scape, 
all of which are creamy white ; clypeal margin with four denticles, the 
medial pair being fused at the base ; hair lobes very broad, extending 
from the lateral denticles to the eyes; clypeal process surface slightly 
convex; mandibles with one prominent denticle; apical antennal 
segments blunt and shghtly curved; antennae amber except for a 
yellow patch on the scape. 

Thorax black except for a divided band on the pronotum and an 
emarginate or divided band on the scutellum, an evanescent patch on 
the metanotum, patches on the propodeum, patch on the pleuron, 
and the tegulae, all of which are creamy white; tegulae low and 
smooth; enclosure smooth except for a prominent medial groove and 
lateral pits; mesosternal tubercles absent; legs variable in color, fore- 
legs hght to dark amber basally to the apical part of femora, beyond 
which they are creamy white; midlegs largely creamy white except 
most of the femora, which are hght to dark amber; hindlegs hght to 
dark amber except the coxae, trochanter, basal part of tibae, and most 
of the tarsi, all of which are creamy white; wings subhyaline, slightly 
clouded toward the apex. 



436 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Abdomen black with subequal creamy-white bands covering about 
one-third of each tergmn; venter black with creamy-white spots 
laterally on 3rd, 4th, and 5th sternites; pygidium oval with the apical 
end trmicate. 

C. femurrubrum Viereck and Cockerell is very different from all 
other species but in several structural characters is close to C. macro- 
sticta Viereck and Cockerell. Both species have been known to use 
Tenebrionidae as prey for their young. 

The males of C. femurrubrum Viereck and Cockerell show little 
variation in color throughout the range of the species; however, the 
females show two distinct color forms, which are recognized here as 
separate subspecies. The reddish form has been found only from the 
Colorado River area east. Since the type male of C. femurrubrum 
Viereck and Cockerell is from Albuquerque, N. Mex., this reddish 
form of female is placed under the subspecies femurrubrum femur- 
rubrum Viereck and Cockerell. The black and yellow form of female 
is taken almost exclusively in southern California. This is the color 
form of the female described as C. athene Banks from Claremont, 
CaUfornia. This is placed under the subspecies C. femurrubrum 
athene Banks. 




Figures 51-53. — Localities of: 51, C. femurrubrum femurrubrum Viereck and Cockerell 
(females only); 52, C. femurrubrum athene Banks (females only); 53, C. femurrubrum 
Viereck and Cockerell (males only). 



Types. — The type male of C. femurrubrum Viereck and Cockerell, 
from Albuquerque, N. Mex., June 30 (Cockerell), is at the Philadelphia 
Academy of Natural Sciences, no. 10040. The type male of C. thione 
Banks, from Colton, Calif. (Pilate), is at the Museum of Comparative 
Zoology, Harvard, no. 23593. 

Distribution. — Southern Nevada, southern Utah, southern Colo- 



WASP GENUS CERCERIS — SCULLEN 437 

rado, and south into Arizona, New Mexico, and southwestern Texas. 

Prey records. — None. 

Plant records. — Acacia sp. (Arizona), A. augustissima (Arizona), 
alfalfa (Arizona), Aplopajypus sp. (Arizona), Asclepias sp. (New 
Mexico), A. subverticillata (Arizona), Baccharis sp. (Arizona), B. 
glutinosa (Arizona), Cissus trifoliata (Arizona), Cleome sp. (Arizona), 
cotton (Ai'izona), Croton sp. (Arizona, New Mexico), Dondm nigra 
(Nevada), Eriogonum sp. (Arizona, New Mexico), E. abertianum 
neomexicanum (Arizona), E. thomasii (Arizona), Gutierrezia sp. 
(Arizona), Haplopappus hartwegi (Arizona), Helianthus sp. (Arizona), 
Lepidium sp. (Arizona), Melilotus alba (Arizona), Mimosa sp. (Ari- 
zona), pigweed leaves (Arizona), Sapindus saponari (Arizona), Wis- 
lizenia sp. (Arizona), W. refracta (Arizona). 

38b. Cerceris feniurrubrum athene Banks, new status 

Figure 52 

Cerceris athene Banks, 1947, pp. 20-21.— Scullen, 1951, p. 1005.— Wasbauer, 1957, 
p. 131.— Scullen, 1960, p. 78. 

Female. — Length 12 mm. Black, yellow, and amber; punctation 
small and somewhat sparse; pubescence average. 

Head slightly wider than the thorax; black except for large frontal 
eye patches on the face, small spots on the lateral clypeal lobes, two 
elongate patches on the vertex, small round patches back cf the eye, 
and base of mandibles, all of which are yellow; most of the clypeus 
and most of the mandibles are dark amber; clypeal margin with a 
broad extension on the medial lobe flanked by a distinct denticle on 
each side and laterad of each of these denticles is another separate 
and much larger denticle; a cone-shaped process on the surface of the 
medial lobe of the clypeus pointing ventrad; mandibles with a single 
basal denticles, apicad of which is a prominent ridge; antennae normal 
in form, amber, becoming darker apically. 

Thorax black except for an emarginate band on the prothorax, the 
scutellmn, the metanotum, very large areas on the propodeum, a 
patch on the pleuron, and the tegulae, all of which are yellow; tegulae 
are low and smooth ; enclosm-e smooth except for a faint medial groove 
and small pits along the lateral sides; mesosternal tubercles absent; 
legs largely amber; wmgs subhyalme but somewhat clouded toward 
the apex. 

Abdomen: all terga yellow emarginate with black anteriorly; 
venter dark amber; pygidium with sides convex, slightly converging 
apically and the apical end truncate. 

Male. — Indistinguishable from the male of Jemurrubrum jemur- 
rubrum Viereck and Cockerell. 



VOL. 116 



438 PROCEEDINGS OF THE NATIONAL MUSEUM 

Types— The type female of C. athene Banks, from Claremont, 
Calif. (C. F. Baker), is at the Museum of Comparative Zoology, 

Harvard, no. 23537. 

Distribution.— Southern California, with a few scattered records 
east of the Colorado River in Arizona, New Mexico, and Texas. 

Prey ubcord. —Eurymetopon rufipes Escholtz (Tenebriomdae) at 
Mecca, Riverside Co., Cahf., July 23, 1956 (Wasbauer, 1957, p. 131). 

Plant recobd. —Asclepias sp. (California), Bacchans sp. (Texas), 
B glutinosa (California), Coyote melon (CaUfornia), Croton calijornicus 
(California), Eremocarpus setigerus (California), Eriogonum fascicula- 
tum (Californ